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.     

Plankton metabolism and bacterial growth efficiency in offshore waters along a latitudinal transect between the UK and Svalbard

Euphotic zone gross primary production, community respiration and net community production were determined from in vitro changes of dissolved oxygen, and from in vivo INT reduction capacity fractionated into two size classes, in offshore waters along a latitudinal transect crossing the North, Norwegian and Greenland Seas between the UK and Svalbard. Rates of gross primary production were higher and more variable than community respiration, so net autotrophy prevailed in the euphotic zone with an average net community production of 164±64 mmol O₂ m⁻² d⁻¹. Respiration seemed to be mainly attributed to large eukaryotic cells (>0.8 µm) with smaller cells, mainly bacteria, accounting for a mean of 25% (range 5–48%) of community respiration. Estimates of bacterial growth efficiency were very variable (range 7–69%) due to uncoupling between bacterial respiration and production. Larger cells tended to contribute more towards total respiration in communities with high gross primary production and low community respiration, while bacteria contributed more towards total respiration in communities with lower gross primary production, typical of microbial-dominated systems. This suggests that community respiration is related to the size structure of the plankton community.     

Phytoplankton dynamics in the NE subarctic Pacific

Ocean Station Papa (OSP, 50°N 145°W) in the NE subarctic Pacific is characterised as high nitrate low chlorophyll (HNLC). However, little is known about the spatial extent of these HNLC waters or the phytoplankton dynamics on the basin scale. Algal biomass, production and size-structure data are presented from winter, spring and summer between 1992 and 1997 for five stations ranging from coastal to open-ocean conditions. The inshore stations (P04–P16) are characterised by the classical seasonal cycle of spring and late summer blooms (production >3 g C m⁻² d⁻¹), diatoms are not Fe-stressed, and growth rate is probably controlled by macronutrient supply. The fate of the phytoplankton is likely sedimentation by diatom-dominated spring blooms, with a pelagic recycling system predominating at other times. The offshore stations (P20/OSP) display low seasonality in biomass and production (OSP, mean winter production 0.3 g C m⁻² d⁻¹, mean spring/summer production 0.85 g C m⁻² d⁻¹), and are dominated by small algal cells. Low Fe availability prevents the occurrence of diatom blooms observed inshore. The main fate of phytoplankton is probably recycling through the microbial food web, with relatively low sedimentation compared to inshore. However, the supply of macro- and micro-nutrients to the coastal and open ocean, respectively, may vary between years. Variability in macro-nutrient supply to the coastal ocean may result in decreased winter reserve nitrate, summer nitrate limitation, subsequent floristic shifts towards small cells, and reduced primary production. Offshore, higher diatom abundances are occasionally observed, perhaps indicating episodic Fe supply. The two distinct oceanic regimes have different phytoplankton dynamics resulting in different seasonality, community structure and fate of algal carbon. These differences will strongly influence the biogeochemical signatures of the coastal and open-oceanic NE subarctic Pacific.     

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.     

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.     

Diapycnal nutrient fluxes on the northern boundary of Cape Ghir upwelling region

In this study we estimate diffusive nutrient fluxes in the northern region of Cape Ghir upwelling system (Northwest Africa) during autumn 2010. The contribution of two co-existing vertical mixing processes (turbulence and salt fingers) is estimated through micro- and fine-structure scale observations. The boundary between coastal upwelling and open ocean waters becomes apparent when nitrate is used as a tracer. Below the mixed layer (56.15±15.56 m), the water column is favorable to the occurrence of a salt finger regime. Vertical eddy diffusivity for salt (K_s) at the reference layer (57.86±8.51 m, CI 95%) was 3×10⁻⁵ (±1.89×10⁻⁹, CI 95%) m² s⁻¹. Average diapycnal fluxes indicate that there was a deficit in phosphate supply to the surface layer (6.61×10⁻⁴ mmol m⁻² d⁻¹), while these fluxes were 0.09 and 0.03 mmol m⁻² d⁻¹ for nitrate and silicate, respectively. There is a need to conduct more studies to obtain accurate estimations of vertical eddy diffusivity and nutrient supply in complex transitional zones, like Cape Ghir. This will provide us with information about salt and nutrients exchange in onshore–offshore zones.     

Latitudinal distribution of microbial plankton abundance,production, and respiration in the Equatorial Atlantic in autumn 2000

Phytoplankton and bacterial abundance, size-fractionated phytoplankton chlorophyll-a (Chl-a) and production together with bacterial production, microbial oxygen production and respiration rates were measured along a transect that crossed the Equatorial Atlantic Ocean (10°N–10°S) in September 2000, as part of the Atlantic Meridional Transect 11 (AMT 11) cruise. From 2°N to 5°S, the equatorial divergence resulted in a shallowing of the pycnocline and the presence of relatively high nitrate (>1 μM) concentrations in surface waters. In contrast, a typical tropical structure (TTS) was found near the ends of the transect. Photic zone integrated ¹⁴C primary production ranged from ∼200 mg C m⁻² d⁻¹ in the TTS region to ∼1300 mg C m⁻² d⁻¹ in the equatorial divergence area. In spite of the relatively high primary production rates measured in the equatorial upwelling region, only a moderate rise in phytoplankton biomass was observed as compared to nearby nutrient-depleted areas (22 vs. 18 mg Chl-a m⁻², respectively). Picophytoplankton were the main contributors (>60%) to both Chl-a biomass and primary production throughout the region. The equatorial upwelling did not alter the phytoplankton size structure typically found in the tropical open ocean, which suggests a strong top-down control of primary producers by zooplankton. However, the impact of nutrient supply on net microbial community metabolism, integrated over the euphotic layer, was evidenced by an average net microbial community production within the equatorial divergence (1130 mg C m⁻² d⁻¹) three-fold larger than net production measured in the TTS region (370 mg C m⁻² d⁻¹). The entire region under study showed net autotrophic community metabolism, since respiration accounted on average for 51% of gross primary production integrated over the euphotic layer.     

Organic matter budget in the Southeast Atlantic continental margin close to the Congo Canyon: In situ measurements of sediment oxygen consumption

A study of organic carbon mineralization from the Congo continental shelf to the abyssal plain through the Congo submarine channel and Angola Margin was undertaken using in situ measurements of sediment oxygen demand as a tracer of benthic carbon recycling. Two measurement techniques were coupled on a single autonomous platform: in situ benthic chambers and microelectrodes, which provided total and diffusive oxygen uptake as well as oxygen microdistributions in porewaters. In addition, sediment trap fluxes, sediment composition (Org-C, Tot-N, CaCO₃, porosity) and radionuclide profiles provided measurements of, respectively input fluxes and burial rate of organic and inorganic compounds.The in situ results show that the oxygen consumption on this margin close to the Congo River is high with values of total oxygen uptake (TOU) of 4±0.6, 3.6±0.5 mmol m⁻² d⁻¹ at 1300 and 3100 m depth, respectively, and between 1.9±0.3 and 2.4±0.2 mmol m⁻² d⁻¹ at 4000 m depth. Diffusive oxygen uptakes (DOU) were 2.8±1.1, 2.3±0.8, 0.8±0.3 and 1.2±0.1 mmol m⁻² d⁻¹, respectively at the same depths. The magnitude of the oxygen demands on the slope is correlated with water depth but is not correlated with the proximity of the submarine channel–levee system, which indicates that cross-slope transport processes are active over the entire margin. Comparison of the vertical flux of organic carbon with its mineralization and burial reveal that this lateral input is very important since the sum of recycling and burial in the sediments is 5–8 times larger than the vertical flux recorded in traps.Transfer of material from the Congo River occurs through turbidity currents channelled in the Congo valley, which are subsequently deposited in the Lobe zone in the Congo fan below 4800 m. Ship board measurements of oxygen profiles indicate large mineralization rates of organic carbon in this zone, which agrees with the high organic carbon content (3%) and the large sedimentation rate (19 mm y⁻¹) found on this site. The Lobe region could receive as high as 19 mol C m⁻² y⁻¹, 1/3 being mineralized and 2/3 being buried and could constitute the largest depocenter of organic carbon in the South Atlantic.     

The supply of nutrients due to vertical turbulent mixing: A study at the Porcupine Abyssal Plain study site in the northeast Atlantic

As part of a multidisciplinary cruise to the Porcupine Abyssal Plain (PAP) study site (49°00′N 16°30′W), in June and July 2006, observations were made of the vertical nitrate flux due to turbulent mixing. Daily profiles of nitrate and turbulent mixing, at the central PAP site, give a mean nitrate flux into the euphotic zone of 0.09 (95% confidence intervals: 0.05–0.16) mmol N m⁻² d⁻¹. This is a factor of 50 lower than the mean observed rate of nitrate uptake within the euphotic zone (5.1±1.3 mmol N m⁻² d⁻¹). By using our direct observations to ‘validate’ a previously published parameterisation for turbulent mixing, we further quantify the variability in the vertical turbulent flux across a roughly 100×100 km region centred on the PAP site, using hydrographic data. The flux is uniformly low (0.08±0.26 mmol N m⁻² d⁻¹, the large standard deviation being due to a strongly non-Gaussian distribution) and is consistent with direct measurements at the central site. It is demonstrated that on an annual basis convective mixing supplies at least 40-fold more nitrate to the euphotic zone than turbulent mixing at this location. Other processes, such as those related with mesoscale phenomena, may also contribute significantly.     

Plankton community and bacterial metabolism in Arctic sea ice leads during summer 2010

Microbial plankton metabolism was examined during summer 2010 in sea ice-influenced waters of the Fram Strait, eastern Arctic Ocean. Rates of gross primary production and community respiration were tightly coupled over a wide range of values (33±3–143±6 and 20±3–126±6 mmol O₂ m^{−2  −1}, respectively) leading to a prevalence of positive net community production. The high variability in community respiration, similar to that of gross primary production, suggests that heterotrophic metabolism may exhibit a significant response to environmental change. Bacterial respiration was assessed at similar time scales to bacterial production measurements, by determining the in vivo INT reduction capacity without pre-filtering the community. Bacteria seem to play a major role in total community respiration, contributing between 5% and 61% of total community respiration, indicating that a high fraction of the organic carbon in Arctic planktonic food webs could flow through these microbes.     

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.     

Primary productivity,differential size fraction and pigment composition responses in two Southern Ocean in situ iron enrichments

Two in situ iron-enrichment experiments were conducted in the Pacific sector of the Southern Ocean during summer 2002 (SOFeX). The “north patch,” established within the Subantarctic Zone (∼56°S), was characterized by high nitrate (∼21 mmol m⁻³) but low silicic acid (2 mmol m⁻³) concentrations. North patch iron enrichment increased chlorophyll (Chl) by 12-fold to 2.1 mg m⁻³ and primary productivity (PP_EU) by 8-fold to 188 mmol C m⁻² d⁻¹. Surprisingly, despite low silicic acid concentrations, diagnostic pigment and size-fraction composition changes indicated an assemblage shift from prymnesiophytes toward diatoms. The “south patch,” poleward of the Southern Boundary of the Antarctic Circumpolar Current (SBACC) (∼66°S), had high concentrations of nitrate (∼27 mmol m⁻³) and silicic acid (64 mmol m⁻³). South patch iron enrichment increased Chl by 9-fold to 3.8 mg m⁻³ and PP_EU 5-fold to 161 mmol C m⁻² d⁻¹ but, notably, did not alter the phytoplankton assemblage from the initial composition of ∼50% diatoms. South patch iron addition also reduced total particulate organic carbon:Chl from ∼300 to 100; enhanced the presence of novel non-photosynthetic, but fluorescent, compounds; and counteracted a decrease in photosynthetic performance as photoperiod decreased. These experiments show unambiguously that in the contemporary, high nitrate Southern Ocean increasing iron supply increases primary productivity, confirming the initial premise of the Martin Iron Hypothesis. However, despite a 5-fold increase in PP_EU under iron-replete conditions in late summer, the effect of iron on annual productivity in the Southern Ocean poleward of the SBACC is limited by seasonal ice coverage and the dark of polar winter.     

A mesoscale eddy driving spatial and temporal heterogeneity in the productivity of the euphotic zone of the northeast Atlantic

In this paper we show how different water masses from a similar geographic region provide an explanation for perturbations in the signal of declining productivity at the Porcupine Abyssal Plain (PAP) study site in the Northeast Atlantic. Furthermore we show that the passage of these different water masses is affected by the filamentary instabilities of a cyclonic eddy just southwest of the PAP site. We describe a high-resolution spatial hydrographic survey conducted with a towed instrument package, complemented by biogeochemical sampling. Maximum rates of primary production of 110 mmol C m^-2 d^-1 seen at the centre of the survey area were associated with the passage of an eddy filament and were enhanced 3 fold relative to far-field conditions (∼36 mmol C m^-2 d^-1). The rotation and stirring influence of the eddy resulted in the sequential passage of 3 distinct water masses past the observation point. This understanding of the lateral stirring around the site enabled us to explain the sharp changes observed in daily primary production rates and other biogeochemical parameters. The spatial survey also revealed a fluorescence maxima associated with the cyclonic eddy that was laterally displaced northwards away from the core, an observation supportive of recent modelling studies.     

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.     

Sulfate reduction in Black Sea sediments: in situ and laboratory radiotracer measurements from the shelf to 2000 m depth

Sulfate reduction rate measurements by the ³⁵SO₄²⁻ core injection method were carried out in situ with a benthic lander, LUISE, and in parallel by shipboard incubations in sediments of the Black Sea. Eight stations were studied along a transect from the Romanian shelf to the deep western anoxic basin. The highest rates measured on an areal basis for the upper 0–15 cm were 1.97 mmol m⁻² d⁻¹ on the shelf and 1.54 mmol m⁻² d⁻¹ at 181 m water depth just below the chemocline. At all stations sulfate reduction rates decreased to values <3 nmol cm⁻³ d⁻¹ below 15 cm depth in the sediment. The importance of sulfate reduction relative to the total mineralization of organic matter was very low, 6%, on the inner shelf, which was paved with mussels, and increased to 47% on the outer shelf at 100 m depth. Where the oxic–anoxic interface of the water column impinged on the sea floor at around 150 m depth, the contribution of sulfate reduction increased from >50% just above the chemocline to 100% just below. In the deep sea, mean sulfate reduction rates were 0.6 mmol m⁻² d⁻¹ corresponding to an organic carbon oxidation of 1.3 mmol m⁻² d⁻¹. This is close to the mean sedimentation rate of organic carbon over the year in the western basin. A comparison with published data on sulfate reduction in Black Sea sediments showed that the present results tend to be higher in shelf sediments and lower in the deep-sea than most other data. Based on the present water column H₂S inventory and the H₂S flux out of the sediment, the calculated turnover time of H₂S below the chemocline is 2100 years.     

Benthic O2 distribution and dynamics in a Mediterranean lagoon (Thau,France): An in situ microelectrode study

Benthic oxygen profiles were acquired using microsensors over two seasonal cycles (December 2001, April and August 2002, January and May 2003) at two stations differently affected by shellfish farming activity in the Thau lagoon (French Mediterranean coast). This study was part of the Microbent-PNEC Program on the study of biogeochemical processes at the sediment–water interface in an eutrophicated environment. We explored seasonal and spatial heterogeneity as well as the biogeochemical drivers of oxygen uptake, such as in situ temperature, bottom water oxygenation and organic matter deposition. O₂ consumption rates were determined by using a transport-reaction model. Maximum rates were reached in August and May and minimum rates in December, April or January. The effect of oyster farming on oxygen fluxes was clearly identified with higher diffusive oxygen uptake in the station inside the oyster parks (C5; 36.8 ± 18.5–87.7 ± 40.8 mmol m⁻² d⁻¹), compared with the station lying outside the oyster parks (C4; 8.6 ± 2.1–30.7 ± 8.3 mmol m⁻² d⁻¹). At C5, the large spatial heterogeneity was statistically concealing temporal variation, whereas a clear statistical difference between cold and warm periods appeared at C4. In these lagoon sediments, the seasonal dynamics of diffusive oxygen demand and consumption rates were mainly driven by seasonal temperature variation at both stations, as well as by seasonal organic matter delivery to the sediment at the station located outside the oyster parks. In the station located below the oyster parks, seasonal variation of organic matter deposition was dampened by oyster filtering activity. Seasonal temperature variation thus appeared as the major driver of oxygen dynamics in this station. Measurements of total O₂ uptake rates indicated a significant fraction of microbial recycling and diffusive transport in oxygen uptake at the station located close to the oyster parks. In the open water site, fauna-mediated O₂ transport prevailed in April 2002 (cold conditions), whereas the microbial recycling seemed to dominate in May 2003 (warm conditions).     

Picophytoplankton dynamics and production in the Arabian Sea during the 1995 Southwest Monsoon

Phytoplankton community structure is expected to shift to larger cells (e.g., diatoms) with monsoonal forcing in the Arabian Sea, but recent studies suggest that small primary producers remain active and important, even in areas strongly influenced by coastal upwelling. To better understand the role of smaller phytoplankton in such systems, we investigated growth and grazing rates of picophytoplankton populations and their contributions to phytoplankton community biomass and primary productivity during the 1995 Southwest Monsoon (August–September). Environmental conditions at six study stations varied broadly from open-ocean oligotrophic to coastal eutrophic, with mixed-layer nitrate and chlorophyll concentrations ranging from 0.01 to 11.5 μM NO₃ and 0.16 to 1.5 μg Chl a. Picophytoplankton comprised up to 92% of phytoplankton carbon at the oceanic stations, 35% in the diatom-dominated coastal zone, and 26% in a declining Phaeocystis bloom. Concurrent in situ dilution and ¹⁴C-uptake experiments gave comparable ranges of community growth rates (0.53–1.05 d⁻¹ and 0.44–1.17 d⁻¹, to the 1% light level), but uncertainties in C:Chl a confounded agreement at individual stations. Microzooplankton grazing utilized 81% of community phytoplankton growth at the oligotrophic stations and 54% at high-nutrient coastal stations. Prochlorococcus (PRO) was present at two oligotrophic stations, where its maximum growth approached 1.4 d⁻¹ (two doublings per day) and depth-integrated growth varied from 0.2 to 0.8 d⁻¹. Synechococcus (SYN) growth ranged from 0.5 to 1.1 d⁻¹ at offshore stations and 0.6 to 0.7 d⁻¹ at coastal sites. Except for the most oligotrophic stations, growth rates of picoeukaryotic algae (PEUK) exceeded PRO and SYN, reaching 1.3 d⁻¹ offshore and decreasing to 0.8 d⁻¹ at the most coastal station. Microzooplankton grazing impact averaged 90, 70, and 86% of growth for PRO, SYN, and PEUK, respectively. Picoplankton as a group accounted for 64% of estimated gross carbon production for all stations, and 50% at high-nutrient, upwelling stations. Prokaryotes (PRO and SYN) contributed disproportionately to production relative to biomass at the most oligotrophic station, while PEUK were more important at the coastal stations. Even during intense monsoonal forcing in the Arabian Sea, picoeukaryotic algae appear to account for a large portion of primary production in the coastal upwelling regions, supporting an active community of protistan grazers and a high rate of carbon cycling in these areas.     

Downward transport of organic carbon by diel migratory micronekton in the western equatorial Pacific:: its quantitative and qualitative importance

Using simultaneous sampling with a commercial-sized trawl, a zooplankton net, and a sediment trap, we evaluated the contribution of vertically migrating micronekton to vertical material transport (biological pump) at two stations (3°00′N, 146°00′E and 3°30′N, 145°20′E) in the western equatorial North Pacific. The gravitational sinking particulate organic carbon flux out of the euphotic zone was 54.8 mg C m⁻² day⁻¹. The downward active carbon flux by diel migrant mesozooplankton was 23.53 and 9.97 mg C m⁻² day⁻¹, and by micronekton 4.40 and 2.26mg C m⁻² day⁻¹ at the two stations. Assuming that the micronekton sampling efficiency of the trawl was 14%, we corrected the downward carbon flux due to micronekton respiration to 29.9 and 15.2mg C m⁻² day⁻¹, or 54.6 and 27.7% of the sinking particle flux at the two stations. The corrected micronekton gut fluxes were 1.53 and 0.97mg C m⁻² day⁻¹. The role of myctophid fish fecal matter as a possible food resource for deep-sea organisms, based on its fatty acid and amino acid analysis, is discussed.     

Zooplankton herbivory in the Arabian Sea during and after the SW monsoon, 1994

Microzooplankton herbivory in the Arabian Sea was measured using dilution experiments towards the end of the SW monsoon in September and during the intermonsoon to NE monsoon period in November–December 1994. Microzooplankton grazing resulted in a turnover of phytoplankton stocks that ranged from 11 to 49% per day. This was equivalent to grazing fluxes of between 1 and 17 mg C m^-3 d^-1. Depth-integrated microzooplankton herbivory ranged between 161 and 415 mg C m^-2 d^-1 during the SW monsoon cruise, and between 110 and 407 mg C m^-2 d^-1 during the intermonsoon period. Microzooplankton grazed between 4 and 60% of daily primary production, with higher percentages found during the intermonsoon season. Phytoplankton growth rates during the SW monsoon ranged from 0.3 to 1.8 d^-1, with lower values in upwelling waters and higher values in downwelling and oligotrophic areas. During the intermonsoon period, phytoplankton growth was more uniform across the basin and averaged 0.68±0.15 d^-1. Microzooplankton abundance in experimental samples varied between 2800 and 16 162 cells l^-1, equivalent to a biomass of between 1.1 and 7.2 mg C m^-3. The mean cell carbon content of microzooplankton was similar in both periods and ranged from 0.33 to 0.55 ng C cell^-1. Microzooplankton were smallest in downwelling waters and largest in oligotrophic waters. Average clearance rates in those taxa that took up fluorescently-labelled algae ranged from 0.2 to 14 μl ind^-1 hr^-1. Average mesozooplankton grazing rates, derived from biomass data, varied from 19 to 92 mg C m^-2 d^-1; these rates accounted for removal of between 4 and 12% of the daily primary production. Mesozooplankton herbivory was most pronounced in upwelling and downwelling waters and reduced in stratified oligotrophic waters during the SW monsoon period. Microzooplankton herbivory was greater than the average mesozooplankton herbivory at all stations, during both the SW monsoon and intermonsoon periods.     

Nitrogen assimilation and the f-ratio in the northwestern Indian Ocean during an intermonsoon period

Rates of nitrogen assimilation by phytoplankton were measured at 13 stations along a transect in the northwestern Indian Ocean, from the Gulf of Oman, southwards to approximately 8°N, during November and December 1994. Nitrate (NO₃), ammonium (NH₄) and urea assimilation were measured using simulated in situ ¹⁵N incubation techniques. These measurements were supported by simultaneous rate measurements of primary production using ¹⁴C incubation techniques and detailed vertical distributions of temperature and chlorophyll concentrations. Euphotic zone integrated nitrogen assimilation rates varied between 1.1 and 23.6 mmol N m^-2 day^-1, with generally higher rates occurring at the northern and southern ends of the transect. At the majority of stations ammonium was the preferred nitrogen substrate assimilated; the average integrated assimilation rate of ammonium being 3.7 mmol N m^-2 day^-1 compared to 1.6 and 1.8 mmol N m^-2 day^-1 for urea and nitrate respectively. This general preference is reflected in the low f-ratios, which were ⩽0.52 for all stations and in the relative preference indices (RPI) values which were consistently >1 for ammonium and <1 for nitrate. A further examination of the data has lead to an apparent partitioning of the northwestern Indian Ocean into 2 regions; a region north of 17°30′N and a region south of this, to about 8°N. This division is based on: (i) the relationship between the f-ratio and ambient nitrate levels; (ii) nitrogen assimilation and primary production and (iii) the biomass distribution. It is suggested that this partitioning should be investigated further with the development of biogeochemical provinces in mind and the estimation of f-ratios on much larger, horizontal scales.