Microbial degradation rates of small peptides and amino acids in the oxygen minimum zone of Chilean coastal waters

We found similar microbial degradation rates of labile dissolved organic matter in oxic and suboxic waters off northern Chile. Rates of peptide hydrolysis and amino acid uptake in unconcentrated water samples were not low in the water column where oxygen concentration was depleted. Hydrolysis rates ranged from 65 to 160 nmol peptide L⁻¹ h⁻¹ in the top 20 m, 8–28 nmol peptide L⁻¹ h⁻¹ between 100 and 300 m (O₂-depleted zone), and 14–19 nmol peptide L⁻¹ h⁻¹ between 600 and 800 m. Dissolved free amino acid uptake rates were 9–26, 3–17, and 6 nmol L⁻¹ h⁻¹ at similar depth intervals. Since these findings are consistent with a model of comparable potential activity of microbes in degrading labile substrates of planktonic origin, we suggest, as do other authors, that differences in decomposition rates with high and low oxygen concentrations may be a matter of substrate lability. The comparison between hydrolysis and uptake rates indicates that microbial peptide hydrolysis occurs at similar or faster rates than amino acid uptake in the water column, and that the hydrolysis of peptides is not a rate-limiting step for the complete remineralization of labile macromolecules. Low O₂ waters process about 10 tons of peptide carbon per h, double the amount processed in surface-oxygenated water. In the oxygen minimum zone, we suggest that the C balance may be affected by the low lability of the dissolved organic matter when this is upwelled to the surface. An important fraction of dissolved organic matter is processed in the oxygen minimum layer, a prominent feature of the coastal ocean in the highly productive Humboldt Current System.     

Microbial respiration in the Levantine Sea: evolution of the oxidative processes in relation to the main Mediterranean water masses

First data on microbial respiration in the Levantine Sea are reported with the aim of assessing the distribution of oxidative processes in association with the main Mediterranean water masses and the changing physical structure determined by the Eastern Mediterranean Transient. Respiratory rates, in terms of metabolic carbon dioxide production, were estimated from measured electron transport system activities in the polygonal area of the Levantine Sea (32.5–36.5 N Latitude, 26.0–30.25 E Longitude) and at Station Geo’95, in the Ionian Sea (35°34.88 N; 17°14.99 E). At the Levantine Sea, the mean carbon dioxide production rate decreased from the upper to the deeper layers and varied from 22.0±12.4 μg C h⁻¹ m⁻³ in the euphotic layer to 1.30±0.5 μg C h⁻¹ m⁻³ in the depth range between 1600 and 3000 m. Significant differences were found among upper, intermediate and bottom layers. The euphotic zone supported a daily carbon dioxide production of 96.6 mg C d⁻¹ m⁻² while the aphotic zone (between 200 and 3000 m) sustained a 177.1 mg C d⁻¹ m⁻² carbon dioxide production. In Station Geo’95, the carbon dioxide production rates amounted to 170.4 and 102.2 mg C d⁻¹ m⁻² in the euphotic and aphotic zones, respectively. The rates determined in the identified water masses showed a tight coupling of respiratory processes and Mediterranean circulation patterns. The increasing respiratory rates in the deep layers of the Levantine Sea are explained by the introduction of younger waters recently formed in the Aegean Sea.     

Community respiration/production and bacterial activity in the upper water column of the central Arctic Ocean

Community metabolism (respiration and production) and bacterial activity were assessed in the upper water column of the central Arctic Ocean during the SHEBA/JOIS ice camp experiment, October 1997–September 1998. In the upper 50 m, decrease in integrated dissolved oxygen (DO) stocks over a period of 124 d in mid-winter suggested a respiration rate of ∼3.3 nM O₂ h⁻¹ and a carbon demand of ∼4.5 gC m⁻². Increase in 0–50 m integrated stocks of DO during summer implied a net community production of ∼20 gC m⁻². Community respiration rates were directly measured via rate of decrease in DO in whole seawater during 72-h dark incubation experiments. Incubation-based respiration rates were on average 3-fold lower during winter (11.0±10.6 nM O₂ h⁻¹) compared to summer (35.3±24.8 nM O₂ h⁻¹). Bacterial heterotrophic activity responded strongly, without noticeable lag, to phytoplankton growth. Rate of leucine incorporation by bacteria (a proxy for protein synthesis and cell growth) increased ∼10-fold, and the cell-specific rate of leucine incorporation ∼5-fold, from winter to summer. Rates of production of bacterial biomass in the upper 50 m were, however, low compared to other oceanic regions, averaging 0.52±0.47 ngC l⁻¹ h⁻¹ during winter and 5.1±3.1 ngC l⁻¹ h⁻¹ during summer. Total carbon demand based on respiration experiments averaged 2.4±2.3 mgC m⁻³ d⁻¹ in winter and 7.8±5.5 mgC m⁻³ d⁻¹ in summer. Estimated bacterial carbon demand based on bacterial productivity and an assumed 10% gross growth efficiency was much lower, averaging about 0.12±0.12 mgC m⁻³ d⁻¹ in winter and 1.3±0.7 mgC m⁻³ d⁻¹ in summer. Our estimates of bacterial activity during summer were an order of magnitude less than rates reported from a summer 1994 study in the central Arctic Ocean, implying significant inter-annual variability of microbial processes in this region.     

Diatom biomass and productivity in oceanic and plume-influenced waters of the western tropical Atlantic ocean

Whereas diatoms (class Bacillariophyceae) often dominate phytoplankton taxa in the Amazon estuary and shelf, their contribution to phytoplankton dynamics and impacts on regional biogeochemistry are poorly understood further offshore in the western tropical Atlantic Ocean (WTAO). Thus, relative contribution of diatoms to phytoplankton biomass and primary production rates and associated environmental conditions were quantified during three month-long cruises in January–February 2001, July–August 2001, and April–May 2003. The upper water column was sampled at 6 light depths (100%, 50%, 25%, 10%, 1% and 0.1% of surface irradiance) at 64 stations between 3° and 14°N latitude and 41° and 58°W longitude. Each station was categorized as ‘oceanic’ or ‘plumewater’, based on principal component analysis of eight physical, chemical and biological variables. All stations were within the North Brazil Current, and plumewater stations were characterized by shallower mixed layers with lower surface salinities and higher dissolved silicon (dSi) concentrations than oceanic stations. The major finding was a much greater role of diatoms in phytoplankton biomass and productivity at plumewater stations relative to oceanic stations. Mean depth-integrated bSi concentrations at the plumewater and oceanic stations were 14.2 and 3.7 mmol m⁻², respectively. Mean depth-integrated SiP rates at the plumewater and oceanic stations were 0.17 and 0.02 mmol m⁻² h⁻¹, respectively. Based on ratios of SiP and PP rates, and typical Si:C ratios, diatoms contributed on average 29% of primary productivity at plumewater stations and only 3% of primary productivity at oceanic stations. In contrast, phytoplankton biomass (as chlorophyll a concentrations) and primary production (PP) rates (as ¹⁴C uptake rates) integrated over the euphotic zone were not significantly different at plumewater and oceanic stations. Chlorophyll a concentrations ranged from 8.5 to 42.4 mg m⁻² and 4.0 to 38.0 mg m⁻² and PP rates ranged from 2.2 to 11.2 mmol m⁻² h⁻² and 1.8 to 10.8 mmol m⁻² h⁻² at plumewater and oceanic stations, respectively. A conservative estimate of annual integrated SiP in offshore waters of Amazon plume between April and August is 0.59 Tmol Si, based on mean SiP rates in plumewaters and satellite-derived estimates of the area of the Amazon plume. In conclusion, river plumewaters dramatically alter the silicon dynamics of the WTAO, forming extensive diatom-dominated phytoplankton blooms that may contribute significantly to the global Si budget as well as contributing to energy and matter flow off of the continental shelf.     

Oxygen distribution and aerobic respiration in the north and south eastern tropical Pacific oxygen minimum zones

Highly sensitive STOX O₂ sensors were used for determination of in situ O₂ distribution in the eastern tropical north and south Pacific oxygen minimum zones (ETN/SP OMZs), as well as for laboratory determination of O₂ uptake rates of water masses at various depths within these OMZs. Oxygen was generally below the detection limit (few nmol L⁻¹) in the core of both OMZs, suggesting the presence of vast volumes of functionally anoxic waters in the eastern Pacific Ocean. Oxygen was often not detectable in the deep secondary chlorophyll maximum found at some locations, but other secondary maxima contained up to ~0.4 µmol L⁻¹. Directly measured respiration rates were high in surface and subsurface oxic layers of the coastal waters, reaching values up to 85 nmol L⁻¹ O₂ h⁻¹. Substantially lower values were found at the depths of the upper oxycline, where values varied from 2 to 33 nmol L⁻¹ O₂ h⁻¹. Where secondary chlorophyll maxima were found the rates were higher than in the oxic water just above. Incubation times longer than 20 h, in the all-glass containers, resulted in highly increased respiration rates. Addition of amino acids to the water from the upper oxycline did not lead to a significant initial rise in respiration rate within the first 20 h, indicating that the measurement of respiration rates in oligotrophic Ocean water may not be severely affected by low levels of organic contamination during sampling. Our measurements indicate that aerobic metabolism proceeds efficiently at extremely low oxygen concentrations with apparent half-saturation concentrations (K_m values) ranging from about 10 to about 200 nmol L⁻¹.     

Dry atmospheric deposition and diazotrophy as sources of new nitrogen to northwestern Mediterranean oligotrophic surface waters

Atmospheric dry deposition of nitrogen (N) and dinitrogen (N₂) fixation rates were assessed in 2004 at the time-series DYFAMED station (northwestern Mediterranean, 43°25′N, 7°52′E). The atmospheric input was monitored over the whole year. Dinitrogen fixation was measured during different seasonal trophic states (from mesotrophy to oligotrophy) sampled during nine cruises. The bioavailability of atmospherically deposited nutrients was estimated by apparent solubility after 96 h. The solubility of dry atmospheric N deposition was highly variable (from ∼18% to more than 96% of total N). New N supplied to surface waters by the dry atmospheric deposition was mainly nitrate (NO₃⁻) (∼57% of total N, compared to ∼6% released as ammonium (NH₄⁺)). The mean bioavailable dry flux of total N was estimated to be ∼112 μmol m⁻² d⁻¹ over the whole year. The NO₃⁻ contribution (70 μmol NO₃⁻ m⁻² d⁻¹) was much higher than the NH₄⁺ contribution (1.2 μmol NH₄⁺ m⁻² d⁻¹). The N:P ratios in the bioavailable fraction of atmospheric inputs (122.5–1340) were always much higher than the Redfield N:P ratio (16). Insoluble N in atmospheric dry deposition (referred to as “organic” and believed to be strongly related to anthropogenic emissions) was ∼40 μmol m⁻² d⁻¹. N₂ fixation rates ranged from 2 to 7.5 nmol L⁻¹ d⁻¹. The highest values were found in August, during the oligotrophic period (7.5 nmol L⁻¹ at 10 m depth), and in April, during the productive period (4 nmol L⁻¹ d⁻¹ at 10 m depth). Daily integrated values of N₂ fixation ranged from 22 to 100 μmol N m⁻² d⁻¹, with a maximum of 245 μmol N m⁻² d⁻¹ in August. No relationship was found between the availability of phosphorus or iron and the observed temporal variability of N₂ fixation rates. The atmospheric dry deposition and N₂ fixation represented 0.5–6% and 1–20% of the total biological nitrogen demand, respectively. Their contribution to new production was more significant: 1–28% and 2–55% for atmospheric dry deposition and N₂ fixation, respectively. The dry atmospheric input was particularly significant in conditions of water column stratification (16–28% of new production), while N₂ fixation reached its highest values in June (46% of new production) and in August (55%).     

New and regenerated production in the Almeria-Oran front area,eastern Alboran Sea

Uptake and regeneration of nitrogen in the Almeria-Oran frontal zone (SW Mediterranean) and adjacent (Atlantic and Mediterranean) systems were studied during the Almofront I cruise (JGOFS-France). The frontal zone was characterized by an upsloping of nitracline from about 50 m in the adjacent systems to 25–30 m within. Along with nitrate, ammonium, chlorophyll a and particulate organic nitrogen also were at higher concentrations in the frontal zone than in the adjacent waters.The nitrate uptake rates were significantly higher in the frontal zone (up to 6.4 nmol l⁻¹ h⁻¹) than in the Atlantic and Mediterranean waters (generally <1 nmol l⁻¹ h⁻¹) indicating a significant increase of new production at the front. This increase was related to the upsloping of the nitracline as shown by the significant correlation (p<0.05) between new production and depth of the nitracline. The new production in the Almeria-Oran was much lower than those recorded in other oceanic and coastal fronts. This could be related to the fact that the nitracline did not rise up to the surface and the high concentrations of nitrate were confined to deeper layers where the ambient light intensity was less. Nitrate uptake in the frontal zone was significantly higher, by 1.7–5.8 times (average 4.2), than the calculated diffusive flux of nitrate, suggesting that vertical advection may be an important source of nitrate. New production rates at the front were also significantly higher (3–9 times, average 5.8) than the PON flux to 100 m depth estimated by sediments traps (Journal of Marine Systems 5, 377–389), suggesting a strong decoupling between surface production and downward flux of POM in the frontal zone.The north–south gradient observed with different parameters indicates the presence of a transfrontal secondary circulation. This distribution also suggests that the primary production in the front is initially nitrate-based, with a diatom-herbivore food chain, whereas regenerated production, associated with an intense recycling of organic matter, later becomes progressively important in time and space.     

Physical drivers of phytoplankton production in the southern Benguela upwelling system

Investigations of primary production (PP) were undertaken in the southern Benguela ecosystem during two research surveys in October 2006 and May 2007. Significant differences in environmental conditions, as well as biomass and PP, were observed between October and May. During October, integrated biomass and PP were significantly higher, ranging from 20.43 to 355.01 mg m⁻², and 0.71 to 6.98 g C m⁻² d⁻¹, respectively, than in May, where the range was 47.92–141.79 mg m⁻², and 0.70–3.35 g C m⁻² d⁻¹, respectively. Distribution patterns indicated low biomass and PP in newly upwelled water along the coast, higher biomass and PP in the mid-shelf region, while lower values were observed at and beyond the shelf edge. Latitudinal variations showed consistently higher biomass and PP in the St. Helena Bay region compared to biomass and PP south of Cape Town. During both surveys, phytoplankton communities were comprised primarily of diatoms and small flagellates, with no significant differences. Phytoplankton adaptation to environmental variability was characterised by increased P_m^B and E_k under elevated temperatures and irradiance, while no clear relationships were evident for α^B. Generalised Additive Models (GAMs) showed that photosynthetic parameters were all significant predictors of photosynthesis rates (P_z), with P_m^B being the most important, accounting for 36.97% of the deviance in P_z. However, biomass levels and environmental conditions exerted a much greater influence on P_z, with irradiance explaining the largest proportion (68.24%) of the deviance. Multiple predictor GAMs revealed that 96.26% of the deviance in P_z could be explained by a model which included nitrate, chlorophyll a, and irradiance.     

Biological versus physical processes as drivers of large oscillations of the air–sea CO2 flux in the Antarctic marginal ice zone during summer

The fugacity of CO₂ and abundance of chlorophyll a (Chla) were determined in two long transects from the Polar Front to the Antarctic Continent in austral summer, December 1995–January 1996. Large undersaturations of CO₂ in the surface water were observed coinciding with high Chla content. In the major hydrographic regions the mean air–sea fluxes were found to range from −3 to +7 mmol m⁻² d⁻¹ making these regions act as a sink as well as a source for CO₂. In the total 40-d period, the summation of the several strong source and sink regions revealed an overall modest net source of 0.3 mmol m⁻² d⁻¹, this based on the Wanninkhof (J. Geophys. Res. 97 (1992) 7373) quadratic relationship at in situ windspeed. A simple budget approach was used to quantify the role of phytoplankton blooms in the inorganic carbonate system of the Antarctic seas in a time frame spanning several weeks. The major controlling physical factors such as air–sea flux, Ekman pumping and upwelling are included. Net community production varies between −9 and +7 mmol m⁻² d⁻¹, because of the large oscillations in the dominance of autotrophic (CO₂ fixation) versus heterotrophic (CO₂ respiration) activity. Here the mixed layer depth is the major controlling factor. When integrated over time the gross influx and efflux of CO₂ from air to sea is large, but the net residual air/sea exchange is a modest efflux from sea to atmosphere.     

Impact of submarine hydrothermal vents on the metal composition of krill and its excretion products

We have measured the metal composition and estimated the excretion rate of trace elements (Ag, Cd, Co, Cu, Ni, Pb, V and Zn) by Antarctic krill (Euphausia superba) in three locations (two located within a submarine hydrothermal vent field and one away from it) along the Antarctic Peninsula region of the Southern Ocean. Results indicated that krill excreted large amounts of Ag, Cu, Pb and Zn, (range: 1.9–41.2 pmol Ag g DW^? 1 h^? 1, 15.3–26.8 nmol Cu g DW^? 1 h^? 1, 308.7–1118.3 pmol Pb g DW^? 1 h^? 1 and 24.4–76.5 nmol Zn g DW^? 1 h^? 1), compared with the non-significant or undetectable release rates of Cd, Co, Ni and V. The metal composition of the excreted material from krill collected in the area of hydrothermal activity was similar to the metal composition reported for suspended particles emitted from those vents. Our results suggest that krill recycling of Ag, Cu, Pb and Zn could potentially influence trace metal concentrations in the water column of the Bransfield region of the Southern Ocean, and that the original source of metals to these waters may be hydrothermal vents.     

Basin-scale variability of phytoplankton biomass,production and growth in the Atlantic Ocean

The latitudinal distributions of phytoplankton biomass, composition and production in the Atlantic Ocean were determined along a 10,000-km transect from 50°N to 50°S in October 1995, May 1996 and October 1996. Highest levels of euphotic layer-integrated chlorophyll a (Chl a) concentration (75–125 mg Chl m⁻²) were found in North Atlantic temperate waters and in the upwelling region off NW Africa, whereas typical Chl a concentrations in oligotrophic waters ranged from 20 to 40 mg Chl m⁻². The estimated concentration of surface phytoplankton carbon (C) biomass was 5–15 mg C m⁻² in the oligotrophic regions and increased over 40 mg C m⁻² in richer areas. The deep chlorophyll maximum did not seem to constitute a biomass or productivity maximum, but resulted mainly from an increase in the Chl a to C ratio and represented a relatively small contribution to total integrated productivity. Primary production rates varied from 50 mg C m⁻² d⁻¹ at the central gyres to 500–1000 mg C m⁻² d⁻¹ in upwelling and higher latitude regions, where faster growth rates (μ) of phytoplankton (>0.5 d⁻¹) were also measured. In oligotrophic waters, microalgal growth was consistently slow [surface μ averaged 0.21±0.02 d⁻¹ (mean±SE)], representing <20% of maximum expected growth. These results argue against the view that the subtropical gyres are characterized by high phytoplankton turnover rates. The latitudinal variations in μ were inversely correlated to the changes in the depth of the nitracline and positively correlated to those of the integrated nitrate concentration, supporting the case for the role of nutrients in controlling the large-scale distribution of phytoplankton growth rates. We observed a large degree of temporal variability in the phytoplankton dynamics in the oligotrophic regions: productivity and growth rates varied in excess of 8-fold, whereas microalgal biomass remained relatively constant. The observed spatial and temporal variability in the biomass specific rate of photosynthesis is at least three times larger than currently assumed in most satellite-based models of global productivity.     

The metabolic balance between autotrophy and heterotrophy in the western Arctic Ocean

The goal of this study was to explore how net community production (NCP) is influenced by the relationship between primary production and community respiration in the western Arctic Ocean. Plankton NCP and respiration were determined by measuring changes in oxygen in light and dark bottle incubations, respectively. Rates of NCP averaged over shelf, slope and basin waters were positive in summer 2002 (57±191 mmol O₂ m⁻² d⁻¹) and spring 2004 (85±86 mmol O₂ m⁻² d⁻¹) and negative in summer 2004 (−25±176 mmol O₂ m⁻² d⁻¹). Determinations of NCP obtained from bottle incubations were similar to rates inferred from in situ changes in dissolved inorganic carbon. An examination of the spatial variability of primary production and community respiration indicated that respiration is distributed more uniformly than primary production. A spatial offset between photosynthesis and respiration from the shelf to the Arctic basin was present in spring 2004, but was not seen at other times. NCP and the potential for export appear to be dependent on an uncoupling of primary production and community respiration. NCP continued into the summer after the stock of NO₃⁻ had been depleted. Our data suggest that the uniform distribution of respiration relative to primary production is an important factor influencing NCP and the potential for export in the western Arctic.     

Dissolved inorganic carbon,alkalinity, nutrient and oxygen seasonal and interannual variations at the Antarctic Ocean JGOFS-KERFIX site

JGOFS-KERFIX (KERguelen point FIXe) time-series station, located south of the polar front in the Indian sector of the Antarctic Ocean, was occupied monthly between January 1990 and March 1995. Annual cycles of dissolved inorganic carbon (DIC), total alkalinity (TALK), oxygen (O₂) and nutrients (nitrate, silicate, phosphate and ammonia) in the upper ocean are presented for this site. From seasonal drawdown of nutrients and DIC, we estimate a spring–summer net community production of 3.2±0.5 mol m⁻² and C/N/P ratios of 100/16/1. The Si/N ratio varies between 1.8 and 3, suggesting low iron concentrations. The spring–summer biogenic silicon export derived from silicate drawdown is 1.18 mol m⁻², consistent with model estimates of silicate export at this site. Seasonal and interannual variations of oxygen, nitrate and DIC due to physical and biological processes are quantified using a simple month-to-month budget formulation. From these budgets, an annual net community production of 5.7±3.3 mol m⁻² yr⁻¹ is estimated, about twice the averaged spring–summer production, indicating that, at KERFIX, there is a positive net community production throughout the year. Air–sea CO₂ fluxes show that KERFIX is a strong CO₂ sink for the atmosphere of 2.4–5.1 mol m⁻² yr⁻¹ in 1993, depending on the gas exchange formulation used. A 2.1–3.3 mol m⁻² yr⁻¹ outgassing of O₂ is observed at KERFIX except in 1993 and 1994 where a decreasing trend of temperature induces an increase of O₂ solubility.     

Photochemical production of the hydroxyl radical in Antarctic waters

Photochemical production rates and steady-state concentrations of the highly reactive OH radical were determined in Antarctic seawater in the Weddell-Scotia Confluence during the austral spring of 1993 and along the Antarctic Peninsula during the austral summer of 1994. OH radical photoproduction rates were 30±2 nM/day and 46±2 nM/day in surface open oceanic and coastal waters, respectively. Corresponding steady-state concentrations were 2.6×10⁻¹⁹ and 4.3×10⁻¹⁹ M which are similar to those found in tropical latitudes. In-situ irradiation experiments (drifter deployments) at different depths in the upper water column indicated that multiple sources for the OH radical existed at three Antarctic stations. Ultrafiltration studies and model calculations based on wavelength-dependent OH radical quantum yields indicated that the main sources were photochemical reactions of low molecular weight dissolved organic matter (DOM), nitrate, and nitrite. Production of the OH radical from nitrate photolysis was almost exclusively UV-B dependent, while OH radical production from nitrite photolysis was mainly UV-A dependent. OH production from DOM photolysis was both UV-A and UV-B dependent. In the upper few meters at open oceanic sites, nitrate and DOM were the dominant OH radical sources, while deeper in the water column DOM and nitrite were important because of the greater importance of UV-A with depth. During non-ozone hole conditions, nitrate contributed about 33%, while DOM plus nitrite contributed about 67% of the predicted OH radical production in open oceanic surface waters. During an ozone hole (151 Dobson units), the corresponding percentages changed to about 40 and 60% for nitrate and DOM due to the higher UV-B irradiance. Model calculations predict that during an ozone hole (151 Dobson units), OH radical production in surface waters will be enhanced by at least 20%, mostly from nitrate photolysis and to a lesser extent from DOM photochemical reactions. This study indicates that ozone hole events significantly increase OH radical production, as well as the photolysis of DOM, in Antarctic waters, and that rates can be as high or higher than those at lower latitudes, especially if differences in temperature and solar irradiance are taken into account.     

Ectoenzymatic activity in surface waters: A transect from the Mediterranean Sea across the Indian Ocean to Australia

The activities of two hydrolytic enzymes (leucine aminopeptidase and β glucosidase), belonging to the particle-bound enzymatic fraction, were measured in open-sea surface waters. Samples were collected along a transect crossing the Indian Ocean during the early NW monsoon period (November and December 2001). The latitudinal pattern of the ectoenzymatic activities highlighted a generally increasing trend of glycolysis approaching the equator, with significantly higher β glucosidase activity (0.79–3.00 nmol l⁻¹ h⁻¹) within the latitudinal range from 12°N to 16°S. In this area, the surface waters coming from the Indonesian Throughflow and the Bay of Bengal carry a considerable quantity of carbohydrates (38.9–41.9 μg l⁻¹), which stimulated glycolytic activity and its cell-specific rates scaled to bacterial abundance. On the other hand, in the Central Indian Ocean, the proteolytic activity was considerable (0.91–2.03 nmol l⁻¹ h⁻¹), although the particulate proteins did not show significant increases and the dissolved proteinlike signal was one of the lowest of the entire transect (0.7 mg l⁻¹ on average compared to the 1.4–1.6 mg l⁻¹ of the adjacent areas). Therefore, in this area, the two ectoenzymes studied did not respond to the same stimulatory effect (namely the specific substrate concentrations). The time needed for the hydrolysis of macromolecules within the particulate and dissolved organic substrate fractions, although these measures are affected by a number of assumptions starting with the potential nature of the ectoenzymatic determinations, confirms these observations. The Central Indian Ocean displayed the lowest values, from 8 to 26 days for particulate and dissolved organic carbon, respectively. As observed in the equatorial areas of the Atlantic Ocean, the relevant degradation activity of the central area of the Indian Ocean Basin suggests a notable heterotrophy based on a faster turnover of organic substrates.     

Winter and spring phytoplankton composition and production in a shallow eutrophic Baltic lagoon

Taxonomic composition and productivity of winter and spring phytoplankton in a eutrophic estuary have been investigated in order to elucidate the carbon flux under conditions of limitation by physical factors – light and temperature. In spite of the important differences in nutrients, solar radiation and water temperature between winter and spring season, mean concentrations of particulate organic carbon were equal to 13.2 and 13.0 mgC l⁻¹, respectively. Chlorophyll a averaged at 79 μgChl l⁻¹ in winter, that is 69% of spring. Although community respiration accounted for only 6–26% of light saturated photosynthesis, integrated net primary production of the 1.2 m deep water column was negative until April. High attenuation of the water body (K_o = 2.9 m⁻¹) lead to a negative carbon balance (net heterotrophy) below 35 cm for all sampling dates. Thus, the high winter POC and phytoplankton values can only originate from summer or autumn primary production. This assumption was supported by a carbon loss rate of just 3% of total organic carbon per day for the whole water column. The composition of phytoplankton was very constant through both seasons: 39% Chlorophyceae, 33% Cyanobacteria and 25% Bacillariophyceae. As expected, phytoplankton was low light acclimated, having high α values (slope of light limited photosynthesis), but moderate maximum photosynthesis rates at saturating irradiances, which were heavily affected by temperature. Calculation of net carbon flux yet showed net heterotrophy of the Bodden waters in winter and early spring were caused by external physical limitation (low surface irradiance and low temperature) in combination with a high light attenuation of the water body.     

Organic carbon flux and remineralization in surface sediments from the northern North Atlantic derived from pore-water oxygen microprofiles

Organic carbon fluxes through the sediment/water interface in the high-latitude North Atlantic were calculated from oxygen microprofiles. A wire-operated in situ oxygen bottom profiler was deployed, and oxygen profiles were also measured onboard (ex situ). Diffusive oxygen fluxes, obtained by fitting exponential functions to the oxygen profiles, were translated into organic carbon fluxes and organic carbon degradation rates. The mean C_org input to the abyssal plain sediments of the Norwegian and Greenland Seas was found to be 1.9 mg C m⁻² d⁻¹. Typical values at the seasonally ice-covered East Greenland continental margin are between 1.3 and 10.9 mg C m⁻² d⁻¹ (mean 3.7 mg C m⁻² d⁻¹), whereas fluxes on the East Greenland shelf are considerably higher, 9.1–22.5 mg C m⁻² d⁻¹. On the Norwegian continental slope C_org fluxes of 3.3–13.9 mg C m⁻² d⁻¹ (mean 6.5 mg C m⁻² d⁻¹) were found. Fluxes are considerably higher here compared to stations on the East Greenland slope at similar water depths. By repeated occupation of three sites off southern Norway in 1997 the temporal variability of diffusive O₂ fluxes was found to be quite low. The seasonal signal of primary and export production from the upper water column appears to be strongly damped at the seafloor. Degradation rates of 0.004–1.1 mg C cm⁻³ a⁻¹ at the sediment surface were calculated from the oxygen profiles. First-order degradation constants, obtained from C_org degradation rates and sediment organic carbon content, are in the range 0.03–0.6 a⁻¹. Thus, the corresponding mean lifetime of organic carbon lies between 1.7 and 33.2 years, which also suggests that seasonal variations in C_org flux are small. The data presented here characterize the Norwegian and Greenland Seas as oligotrophic and relatively low organic carbon deep-sea environments.     

Biogenic fluxes in the Cariaco Basin: a combined study of sinking particulates and underlying sediments

The fluxes of total mass, organic carbon (OC), biogenic opal, calcite (CaCO₃) and long-chain C₃₇ alkenones (ΣAlk₃₇) were measured at three water depths (275, 455 and 930 m) in the Cariaco Basin (Venezuela) over three separate annual upwelling cycles (1996–1999) as part of the CARIACO sediment trap time-series. The strength and timing of both the primary and secondary upwelling events in the Cariaco Basin varied significantly during the study period, directly affecting the rates of primary productivity (PP) and the vertical transport of biogenic materials. OC fluxes showed a weak positive correlation (r²=0.3) with PP rates throughout the 3 years of the study. The fluxes of opal, CaCO₃ and ΣAlk₃₇ were strongly correlated (0.6<r²<0.8) with those of OC. The major exception was the lower than expected ΣAlk₃₇ fluxes measured during periods of strong upwelling. All sediment trap fluxes were significantly attenuated with depth, consistent with marked losses during vertical transport. Annually, strong upwelling conditions, such as those observed during 1996–1997, led to elevated opal fluxes (e.g., 35 g m⁻² yr⁻¹ at 275 m) and diminished ΣAlk₃₇ fluxes (e.g., 5 mg m⁻² yr⁻¹ at 275 m). The opposite trends were evident during the year of weakest upwelling (1998–1999), indicating that diatom and haptophyte productivity in the Cariaco Basin are inversely correlated depending on upwelling conditions.The analyses of the Cariaco Basin sediments collected via a gravity core showed that the rates of OC and opal burial (10–12 g m⁻² yr⁻¹) over the past 5500 years were generally similar to the average annual water column fluxes measured in the deeper traps (10–14 g m⁻² yr⁻¹) over the 1996–1999 study period. CaCO₃ burial fluxes (30–40 g m⁻² yr⁻¹), on the other hand, were considerably higher than the fluxes measured in the deep traps (∼10 g m⁻² yr⁻¹) but comparable to those obtained from the shallowest trap (i.e. 38 g m⁻² yr⁻¹ at 275 m). In contrast, the burial rates of ΣAlk₃₇ (0.4–1 mg m⁻² yr⁻¹) in Cariaco sediments were significantly lower than the water column fluxes measured at all depths (4–6 mg m⁻² yr⁻¹), indicating the large attenuation in the flux of these compounds at the sediment–water interface. The major trend throughout the core was the general decrease in all biogenic fluxes with depth, most likely due to post-depositional in situ degradation. The major exception was the relatively low opal fluxes (∼5 g m⁻² yr⁻¹) and elevated ΣAlk₃₇ fluxes (∼2 mg m⁻² yr⁻¹) measured in the sedimentary interval corresponding to 1600–2000 yr BP. Such compositions are consistent with a period of low diatom and high haptophyte productivity, which based on the trends observed from the sediment traps, is indicative of low upwelling conditions relative to the modern day.     

Commercially-cultured oysters (Crassostrea gigas) exert top-down control on intertidal pelagic resources in Willapa Bay,Washington, USA

The capacity of filter feeders to reduce seston and phytoplankton concentrations in the water column has important implications for restoration and management of coastal ecosystems. We directly measured changes in chlorophyll a concentration on commercially stocked intertidal oyster beds (Crassostrea gigas) in Willapa Bay, Washington, USA by recording water properties near small drifters as they tracked parcels of water across tide flats. Chlorophyll declined 9.6% per half hour in water passing on-bottom adult oysters and 41% for longline adult oysters, whereas chlorophyll concentrations increased as water flowed across tide flats without adult oysters. Field filtration rates, which were fit to exponential declines in chlorophyll and accounted for oyster density and water depth, averaged 0.35 L g^− 1 h^− 1 (shucked dry weight) for on-bottom aquaculture and 0.73 L g^− 1 h^− 1 for longline culture, compared to values of 2.5–12 L g^− 1 h^− 1 reported from laboratory studies of C. gigas. Field filtration rates may be lower than laboratory rates due to unfavorable field conditions (e.g., low initial chlorophyll concentrations) or masked by resuspension of benthic microalgae. In addition to distinctions among on-bottom, longline, and no-oyster habitats, Akaike's Information Criterion analysis showed temperature, initial chlorophyll concentration, and depth related to chlorophyll decline. This research corroborates mathematical models suggesting that benthic suspension feeders are exerting top-down control of pelagic production in this estuary, with strong patterns in chlorophyll emerging across extensive tideflats populated by C. gigas despite low field filtration rates.     

Effects of solar radiation on dimethylsulfide cycling in the western Atlantic Ocean

The influence of solar radiation on springtime rates of photochemical and biological consumption of dimethylsulfide (DMS) in surface waters from the western Atlantic Ocean was examined by exposing 0.2 μm filtered and unfiltered surface seawater to natural sunlight at five depths in the upper 30 m. Parallel deck incubations of 0.2 μm filtered seawater under various long-pass optical filters were also carried out to aid in assessing the wavelength dependence of DMS photolysis. DMS photolysis rate constants for mid-day exposure (∼10:30–17:30 local time) to surface irradiance ranged from 0.026 to 0.086 h⁻¹ and were highest in coastal and shelf waters. Photolysis rate constants decreased with increasing irradiation depth, in accordance with the attenuation of ultraviolet radiation (UVR, 280–400 nm). Total DMS consumption rates (photochemical+biological) in unfiltered surface samples also decreased with increasing incubation depth and were larger than photolysis rates at nearly all depths and all stations. The decrease in photolysis rate constants with exposure depth was mirrored by biological DMS consumption rate constants that were severely inhibited at surface irradiances, and approached or exceeded dark rate constants at deeper exposure depths. Photolysis rates were 2–19 times greater than estimated biological consumption rates in the surface light exposed samples, while biological consumption rates were significantly larger than photolysis rates at incubation depths below the 1% light level for UV–B radiation (280–320 nm). Total DMS loss rates increased up to nine-fold with UVR exposure, but changes in DMS concentrations were not strongly correlated to light dose, presumably due to parallel, light-mediated DMS production processes. The primary loss process for DMS depended mainly on the depth interval considered and the attenuation of UVR; in general, photochemical removal dominated shallow layers characterized by high UV–B intensities, whereas biological removal dominated in deeper layers where UV–B was absent, but UV–A (320–400 nm) and visible (400–700 nm) light fluxes were still relatively high. These results demonstrate that UVR exposure significantly influences the spatial and temporal pattern of DMS production and loss processes, and ultimately the DMS flux to the atmosphere.