Mesozooplankton in the Arctic Ocean in summer

The biomass, species and chemical composition of the mesozooplankton and their impact on lower food levels were estimated along a transect across the Arctic Ocean. Mesozooplankton biomass in the upper 200 m of the water column was significantly higher (19–42 mg DW m^-3) than has previously been reported for the Arctic Ocean, and it reached a maximum at ca. 87°N in the Amundsen Basin. The lowest values were recorded in the Chukchi Sea and Nansen Basin, where ice cover was lower (50–80%) than in the central Arctic Ocean. In the deeper strata (200–500 m) of the Canadian and Eurasian Basins, the biomass was always much lower (4.35–16.44 mg DW m^-3). The C/N (g/g) ratio for the mesozooplankton population was high (6.5–8.5) but within the documented range. These high values (when compared to 4.5 at lower latitudes) may be explained by the high lipid content. Mesozooplankton accounted for approximately 40% of the total particulate organic carbon in the upper 100 m of the water column. Mesozooplankton species composition was homogeneous along the transect, consisting mainly of copepods (70–90% of the total number). It was dominated by four large copepod species (Calanus hyperboreus, C. glacialis, C. finmarchicus and Metridia longa), which together accounted for more than 80% of the total biomass. According to measurements of gut pigment and gut turnover rates, the mesozooplankton on average ingested between 6 and 30% of their body carbon per day as phytoplankton. Microzooplankton may have provided an additional source of energy for the mesozooplankton community. These data emphasize the importance of mesozooplankton in the arctic food web and reinforce the idea that the Arctic Ocean should no longer be considered to be a “biological desert”.     

Regional variations in the fluxes of foraminifera carbonate,coccolithophorid carbonate and biogenic opal in the northern Indian Ocean

Mass fluxes of diatom opal, planktonic foraminifera carbonate and coccolithophorid carbonate were measured with time-series sediment traps at six sites in the Arabian Sea, Bay of Bengal and Equatorial Indian Ocean (EIOT). The above fluxes were related to regional variations in salinity, temperature and nutrient distribution. Annual fluxes of diatom opal range between 3 and 28 g m⁻² yr⁻¹, while planktonic foraminifera carbonate fluxes range between 6 and 23 g m⁻² yr⁻¹ and coccolithophorid carbonate fluxes range between 4 and 24 g m⁻² yr⁻¹. Annual planktonic foraminifera carbonate to coccolithophorid carbonate ratios range between 0.8 and 2.2 and coccolithophorid carbonate to diatom opal ratios range between 0.5 and 3.3.In the western Arabian Sea, coccolithophorids are the major contributors to biogenic flux during periods of low nutrient concentrations. Coccolithophorid carbonate fluxes decrease and planktonic foraminiferal carbonate and diatom opal fluxes increase when nutrient-rich upwelled waters are advected over the trap site. In the oligotropic eastern Arabian Sea, coccolithophorid carbonate fluxes are high throughout the year. Planktonic foraminiferal carbonate fluxes are the major contributors to biogenic flux in the EIOT. In the northern and central Bay of Bengal, when surface salinity values drop sharply during the SW monsoon, there is a drastic reduction in planktonic foraminiferal carbonate fluxes, but coccolithophorid carbonate and diatom opal fluxes remain steady or continue to increase. Distinctly higher annual molar Si_bio/C_inorg (>1) and C_org/C_inorg (>1.5) ratios are observed in the northern and central Bay of Bengal mainly due to lower foraminiferal carbonate production as a result of sharp salinity variations. We can thus infer that the enhanced freshwater supply from rivers should increase oceanic CO₂ uptake. Its silicate supply favours the production of diatoms while the salinity drop produces conditions unfavourable for most planktonic foraminifera species.     

Nitrogen sources for new production in the NE Arabian Sea

New productivity measurements using the ¹⁵N tracer technique were conducted in the north-eastern (NE) Arabian Sea during six expeditions from 2003 to 2007, mostly in winter. Our results indicate that the NE Arabian Sea has a potential for higher new productivity during blooms. Nitrate uptake by plankton is the highest during late winter. New productivity and f-ratios in the NE Arabian Sea are mainly controlled by hydrodynamic and meteorological parameters such as wind strength, sea surface temperature (SST), mixed layer depth (MLD) and mixed layer nitrate. Deepening of the mixed layer supplies nitrate from below, which supports the observed nitrogen uptake. Higher f-ratios during blooms indicate the strong coupling between surface layers and sub-surface layers. Deepening of mixed layer below 100 m (from its inter-monsoon value between 30 and 40 m) transferred often more than 100 mmol N–NO₃ m^? 2 into the surface layers from below. The observed winter blooms in the region are supported by such input and are sustained for more than a month. Higher new productivity has been found in late winter, whereas transport of nitrate is maximum in early winter. In general, new production varies progressively during winter. Diurnal cycling of the mixed layer could be the reason for the under utilization of entrained nitrate during early winter. New productivity values and wind strength show significant differences during Feb–Mar 03 and Feb–Mar 04. These differences indicate that the winter cooling and parameters related the biological productivity also vary inter-annually. However, the difference between the new productivity values between Feb–Mar 03 and Feb–Mar 04 is much lower than the difference between Jan 03 and Feb–Mar 03. The results suggest that amplitude of seasonal variation is higher than the inter-annual variation in the region. During spring, Fickian diffusive fluxes of nitrate into the surface layer range from 0.51 to 1.38 mmol N–NO₃ m^? 2 day^? 1, and can account for 67% and 78% of the observed nitrogen uptake in the coastal and open ocean regions, respectively. We document the intra-seasonal and inter-annual variations in new productivity during winter and identify sources of nitrate which support the observed productivity during spring.     

Monsoonal influence on the distribution of phytoplankton pigments in the Arabian Sea

Variations in the distribution of chemotaxonomic pigments were monitored in the Arabian Sea and the Gulf of Oman at the end of the SW monsoon in September 1994 and during the inter-monsoon period in November/December 1994 to determine the seasonal changes in phytoplankton composition. The Gulf of Oman was characterized by sub-surface chlorophyll maxima at 20-40 m during both seasons, and low levels of divinyl chlorophyll a indicated that prochlorophytes did not contribute significantly to the total chlorophyll a. Prymnesiophytes (19′-hexanoyloxyfucoxanthin), diatoms (fucoxanthin) and chlorophyll b containing organisms accounted for most of the phytoplankton biomass in September, while prymnesiophytes dominated in November/December. In the Arabian Sea in September, high total chlorophyll a concentrations up to 1742 ng l^-1 were measured in the coastal upwelling region and a progressive decline was monitored along the 1670 km offshore transect to oligotrophic waters at 8°N. Divinyl chlorophyll a was not detected along this transect except at the two most southerly stations where prochlorophytes were estimated to contribute 25–30% to the total chlorophyll a. Inshore, the dominance of fucoxanthin and/or hexanoyloxyfucoxanthin indicated that diatoms and prymnesiophytes generally dominated the patchy phytoplankton community, with zeaxanthin-containing Synechococcus also being important, especially in surface waters. At the southern oligotrophic localities, Synechococcus and prochlorophytes dominated the upper 40 m and prymnesiophytes were the most prominent at the deep chlorophyll maximum. During the inter-monsoon season, total chlorophyll a concentrations were generally half those measured in September and highest levels were found on the shelf (1170 ng l^-1). Divinyl chlorophyll a was detected at all stations along the Arabian Sea transect, and we estimated that prochlorophytes contributed between 3 and 28% to the total chlorophyll a, while at the two oligotrophic stations this proportion increased to 51–52%. While procaryotes were more important in November/December than September, eucaryotes still accounted for >50% of the total chlorophyll a. Pigment/total chlorophyll a ratios indicated that 19′-hexanoyloxyfucoxanthin-containing prymnesiophytes were the dominant group, although procaryotes accounted for 65% at the two southerly oligotrophic stations.     

The structure of zooplankton communities,in the 2 to 2000 μm size range,in the Arabian Sea during and after the SW monsoon, 1994

Zooplankton communities, studied in the surface mixed layer on a 1600 m transect across the Arabian Sea, were found to differ in their temporal and spatial response to seasonal forcing. The transect studied, spanned seasonally eutrophic upwelling, mesotrophic downwelling and aseasonal oligotrophic waters. The nano- and microzooplankton communities constituted a relatively constant compartment in the tropical monsoon ecosystem, whilst the mesozooplankton showed a clear response to both upwelling and season. The heterotrophic nanoflagellates were concentrated in the surface mixed layer, except in the eutrophic upwelling waters of the SW monsoon. They reached maximum cell concentrations of 855 ml^-1 during the SW monsoon and a maximum biomass of 8.4 mg C m^-3 during the intermonsoon. Nanozooplankton standing stocks, in the surface mixed layer, ranged between 7 and 333 mg C m^-2, with highest stocks found during the intermonsoon. The microzooplankton community was dominated by Protozoa, particularly aloricate ciliates and heterotrophic dinoflagellates, which accounted for up to 99% in terms of numbers and up to 71% of the biomass. Sarcodines and metazoan nauplii were recorded in lower numbers (<400 l^-1). The microzooplankton were also concentrated in the surface mixed layer during both periods, except in the eutrophic coastal waters during the SW monsoon, when relatively high biomass values were found below the mixed layer depth. Their standing stocks, in the surface mixed layer, ranged between 50 and 182 mg C m^-2, with the highest concentration found in the mesotrophic offshore waters during the late monsoon period. Total mesozooplankton standing stocks, in the surface 100 m, decreased with distance from the coastal to offshore waters and between seasons, decreasing from 1248 to 238 mg C m^-2 during the late SW monsoon and 656–89 mg C m^-2 during the following intermonsoon. The largest size class, of 1000–2000 μm sized organisms, dominated throughout except at the oligotrophic station during the intermonsoon period, when the smallest class, of 200–500 μm, were more important. The shift in size structure from large to small zooplankton occurred in response to a shift in dominance from large to small phytoplankton cells both spatially, along a eutrophic–oligotrophic gradient, and seasonally. These responses are a result of the physical forcing associated with the monsoon seasons in the Arabian Sea.     

Nitrogen dynamics during the Arabian Sea Northeast Monsoon

This investigation focused on the weaker and less well understood of the two Arabian Sea monsoonal wind phases, the NE Monsoon, which persists for 3–4 months in the October to February period. Historically, this period has been characterized as a time of very low nutrient availability and low biological production. As part of the US JGOFS Arabian Sea Process Study, 17 stations were sampled on a cruise in January 1995 (late NE Monsoon) and, 15 stations were sampled on a cruise in November 1995 (early NE Monsoon). Only the southern most stations (10° and 12°N) and one shallow coastal station were as nutrient-depleted as had been expected from the few relevant prior studies in this region. Experiments were conducted to ascertain the relative importance of different nitrogenous nutrients and the sufficiency of local regeneration processes in supplying nitrogenous nutrients utilized in primary production. Except for the southern oligotrophic stations, the euphotic zone concentrations of NO₃⁻ were typically 5–10-fold greater than those of NO₂⁻ and NH₄⁺. There was considerable variation (20–40-fold) in nutrient concentration both within and between the two sections on each cruise. All nitrogenous nutrients were more abundant (2–4-fold) later in the NE Monsoon. Strong vertical gradients in euphotic zone NH₄⁺ concentration, with higher concentrations at depth, were common. This was in contrast to the nearly uniform euphotic zone concentrations for both NO₃⁻ and NO₂⁻. Half-saturation constants for uptake were higher for NO₃⁻ (1.7 μmol kg⁻¹ (s.d.=0.88, n=8)) than for NH₄⁺ (0.47 μmol kg⁻¹ (s.d.=0.33, n=5)). Evidence for the suppressing effect of NH₄⁺ on NO₃⁻ uptake was widespread, although not as severe as has been noted for some other regions. Both the degree of sensitivity of NO₃⁻ uptake to NH₄⁺ concentration and the half-saturation constant for NO₃⁻ uptake were correlated with ambient NO₃⁻ concentration. The combined effect of high affinity for low concentrations of NH₄⁺ and the effect of NH₄⁺ concentration on NO₃⁻ uptake resulted in similarly low f-ratios, 0.15 (s.d.=0.07, n=15) and 0.13 (s.d.=0.08, n=17), for early and late observations in the NE Monsoon, respectively. Stations with high f-ratios had the lowest euphotic zone NH₄⁺ concentrations, and these stations were either very near shore or far from shore in the most oligotrophic waters. At several stations, particularly early in the NE Monsoon, the utilization rates for NO₂⁻ were equal to or greater than 50% the utilization rates for NO₃⁻. When converted with a Redfield C : N value of 6.7, the total N uptake rates measured in this study were commensurate with measurements of C productivity. While nutrient concentrations at some stations approached levels low enough to limit phytoplankton growth, light was shown to be very important in regulating N uptake at all stations in this study. Diel periodicity was observed for uptake of all nitrogenous nutrients at all stations. The amplitude of this periodicity was positively correlated with nutrient concentration. The strongest of these relationships occurred with NO₃⁻. Ammonium concentration strongly influenced the vertical profiles for NO₃⁻ uptake as well as for NH₄⁺ uptake. Both NO₂⁻ and NH₄⁺ were regenerated within the euphotic zone at rates comparable to rates of uptake of these nutrients, and thus maintenance of mixed layer concentrations did not require diffusive or advective fluxes from other sources. Observed turnover times for NH₄⁺ were typically less than one day. Rapid turnover and the strong light regulation of NH₄⁺ uptake allowed the development and maintenance of vertical structure in NH₄⁺ concentration within the euphotic zone. In spite of the strong positive effect of light on NO₂⁻ uptake and its strong negative effect on NO₂⁻ production, the combined effects of much longer turnover times for this nutrient and mixed layer dynamics resulted in nearly uniform NO₂⁻ concentrations within the euphotic zone. Responses of the NE Monsoon planktonic community to light and nutrients, in conjunction with mixed layer dynamics, allowed for efficient recycling of N within the mixed layer. As the NE Monsoon evolved and the mixed layer deepened convectively, NO₂⁻ and NO₃⁻ concentrations increased correspondingly with the entrainment of deeper water. Planktonic N productivity increased 2-fold, but without a significant change the new vs. recycled N proportionality. Consequently, NO₃⁻ turnover time increased from about 1 month to greater than 3 months. This reflected the overriding importance of recycling processes in supplying nitrogenous nutrients for primary production throughout the duration of the NE Monsoon. As a result, NO₃⁻ supplied to the euphotic zone during the NE Monsoon is, for the most part, conserved for utilization during the subsequent intermonsoon period.     

The influence of the south-west monsoon upon the nutrient biogeochemistry of the Arabian Sea

Variations in the nutrient concentrations were studied during two cruises to the Arabian Sea. The situation towards the end of the southwest monsoon season (September/October 1994) was compared with the inter-monsoonal season during November and December 1994. Underway surface transects showed the influence of an upwelling system during the first cruise with deep, colder, nutrient-rich water being advected into the surface mixed layer. During the southwesterly monsoon there was an area of coastal Ekman upwelling, bringing colder water (24.2°C) into the surface waters of the coastal margin. Further offshore at about 350 km there was an area of Ekman upwelling, as a result of wind-stress curl, north of the Findlater Jet axis; this area also had cooler surface water (24.6°C). Further offshore (>1000 km) the average surface temperatures increased to >27°C. These waters were oligotrophic with no evidence of the upwelling effects observed further inshore. In the upwelling regions nutrient concentrations in the close inshore coastal zone were elevated (NO₃=18 μmol l^-1, PO₄=1.48 μmol l^-1); higher concentrations also were measured at the region of offshore upwelling off the shelf, with a maximum nitrate concentration of 12.5 μmol l^-1 and a maximum phosphate concentration of 1.2 μmol l^-1. Nitrate and phosphate concentrations decreased with increasing distance offshore to the oligotrophic waters beyond 1400 km, where typical nitrate concentrations were 35.0 nmol l^-1 (0.035 μmol l^-1) in the surface mixed layer. A CTD section from the coastal shelf, to 1650 km offshore to the oligotrophic waters, clearly showed that during the monsoon season, upwelling is one of the major influences upon the nutrient concentrations in the surface waters of the Arabian Sea off the coast of Oman. Productivity of the water column was enhanced to a distance of over 800 km offshore. During the intermonsoon period a stable surface mixed layer was established, with a well-defined thermocline and nitracline. Surface temperature was between 26.8 and 27.4°C for the entire transect from the coast to 1650 km offshore. Nitrate concentrations were typically between 2.0 and 0.4 μmol l^-1 for the transect, to about 1200 km where the waters became oligotrophic, and nitrate concentrations were then typically 8–12 nmol l^-1. Ammonia concentrations for the oligotrophic waters were typically 130 nmol l^-1, and are reported for the first time in the Indian Ocean. The nitrogen/phosphorus (N/P) ratios suggest that phytoplankton production was potentially nitrogen-limited in all the surface waters of the Arabian Sea, with the greatest nitrogen limitation during the intermonsoon period.     

Seasonal variations in the distribution of Fe and Al in the surface waters of the Arabian Sea

Concentrations of dissolved Al and Fe in the surface mixed layer were measured during five cruises of the 1995 US JGOFS Arabian Sea Process Study, Concentrations of both Al and Fe were relatively uniform between January and April, the NE Monsoon and the Spring Intermonsoon period, ranging from 2 to 11 nM Al (mean 5.3 nM) and 0.5 to 2.4 nM Fe (mean 1.0 nM). In July/August, after the onset of the SW Monsoon, surface water Al and Fe concentrations increased significantly (Al range 4.5–20.1 nM; mean=10 nM, Fe range 0.57–2.4 nM; mean=1.3 nM), particularly in the NE part of the Arabian Sea, as the result of the input and partial dissolution of eolian dust. Using the enrichment of Al in the surface waters, we estimate this is the equivalent to the deposition of 2.2–7.4 g m⁻² dust, which is comparable to values previously estimated for this region. Approximately one month later (August/September), surface water concentrations of both Al and Fe were found to have decreased significantly (mean Al 7.4 nM, mean Fe 0.90 nM) particularly in the same NE region, as the result of export of particulate material from the euphotic zone. Fe supply to the surface waters is also affected by upwelling of sub-surface waters in the coastal region of the Arabian Sea during the SW Monsoon. Despite the proximity of high concentrations of Fe in the shallow sub-oxic layer, freshly upwelled water is not drawn from this layer and the NO₃/Fe ratio in the initially upwelled water is below the value at which Fe limitation is through to occur. Continued deposition of eolian Fe into the upwelled water as it advects offshore provides the Fe required to raise this ratio above the Fe limitation value.     

Seasonal and spatial variability in N2O distribution in the Arabian Sea

Extensive measurements of nitrous oxide (N₂O) were made in the central and eastern Arabian Sea during the northeast monsoon (February–March), intermonsoon (April–May) and southwest monsoon (July–August) seasons. The latitudinal and longitudinal variations, along with seasonal changes with respect to winter convection and coastal upwelling, are clearly discernible. Vertical profiles collected down to 1000 m show that the Arabian Sea water column is supersaturated with N₂O at all depths. However, N₂O consumption at intermediate depths, coincident with the oxygen minimum and associated with sediment–water interfaces, and in the denitrifying zone, coincident with NO^-₂ secondary maxima, are also apparent. The N₂O concentration varies from ∼10 nM near the surface to about 80 nM in the secondary peak region (≈800 m). Interrelationships with chemical parameters suggest nitrification to be the main process for the production of N₂O in the oceanic water. Plots of apparent oxygen utilization vs production of N₂O indicate a consistent linear relationship for AOU between 0 and 200 μM.     

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.     

Bacterioplankton activity in the surface waters of the Arabian Sea during and after the 1994 SW monsoon

Bacterial biomass and production were measured on two cruises to the northwestern Arabian Sea in 1994; the first cruise took place towards the end of the SW monsoon in September, and the second cruise during the inter-monsoon period in November and December. Although phytoplankton production was significantly higher during the monsoon, bacterial numbers showed little difference. Bacteria were most abundant in the euphotic zone and highest bacterial numbers were measured during the monsoon period in the Gulf of Oman and the shelf waters off southern Oman; in these regions, numbers ranged from 0.9 to 1.6×10⁹ bacteria l^-1. On both cruises, bacteria were less abundant in the euphotic zone of the central Arabian Sea and typically ca 0.8×10⁹ cells l^-1 were present. The majority of bacteria (80–95%) were small cocci that were larger (median diameter 0.40 μm) during the monsoon period than the inter-monsoon, when the cells had a diameter of 0.36 μm; there was no comparable change in cell dimensions of bacteria present as rods. Bacterial production was measured by the incorporation of ³H-thymidine and ³H-leucine. On both cruises, uptake rates were highest on the Omani shelf and decreased offshore. In the central Arabian Sea, thymidine incorporation rates were similar in the monsoon and inter-monsoon periods, but higher rates of leucine incorporation were measured during the monsoon period. Bacterial production was a relatively small proportion of phytoplankton production in both periods sampled; bacterial production was equivalent to between 10 and 30% of the daily primary production in the Arabian Sea.     

Mesozooplankton grazing manipulations during in vitro iron enrichment studies in the NE subarctic Pacific

IronEx I demonstrated a rapid and marked response by grazers to Fe-induced increases in phytoplankton stocks, which was thought to be due, in part, to arrested vertical migration by mesozooplankton. These observations prompted an investigation of the relative roles of Fe enrichment and grazing pressure in controlling the magnitude of phytoplankton stocks in the NE subarctic Pacific. The grazing impact of increased mesozooplankton abundance in response to a localised Fe-induced enhancement of algal biomass was simulated by performing in vitro (6 d) grazer perturbation experiments in May 1994 and September 1995 at Ocean Station Papa (OSP), when pelagic mesozooplankton stocks are usually at their annual maximum and submaximal, respectively. Manipulations were designed to increase mesozooplankton stocks in 25L carboys after various lag-times corresponding to grazing pressure greater or equal to that in situ, and to monitor changes in chlorophyll a levels as a proxy for grazing pressure. At the onset of the experiments, in vitro mesozooplankton abundances were comparable to those in situ. Despite the addition of mesozooplankton to selected Fe-enriched carboys in May after 24, 48 and 72 h, corresponding to ca. two-fold increases in their abundances, chlorophyll a increased to ca. 2 μg l⁻¹ in all treatments. In September, chlorophyll a levels increased five-fold to 2 μg l⁻¹ after 4 days – but little thereafter – in the presence of up to ten-fold higher animal abundances (added at t=0) than observed in situ. Thus, Fe-induced increases in diatom growth rates were sufficiently high to escape both initial and additional grazing pressure. If and when Fe is supplied to this region, it is unlikely that mesozooplankton can respond and graze down the resulting elevated algal abundance. Theoretical calculations, based on algal growth and grazing rate data from May in this study, suggested that a greater than five-fold increase in mesozooplankton abundance, after a 48-h lag, is required to exert sufficient grazing pressure to prevent Fe-mediated increases in algal biomass. These findings are discussed in relation to the scale dependency of such events, and the pelagic ecology of other High Nitrate Low Chlorophyll regions.     

Total organic and inorganic carbon exchange through the Strait of Gibraltar in September 1997

The total organic carbon (TOC) and total inorganic carbon (C_T) exchange between the Atlantic Ocean and the Mediterranean Sea was studied in the Strait of Gibraltar in September 1997. Samples were taken at eight stations from western and eastern entrances of the Strait and at the middle of the Strait (Tarifa Narrows). TOC was analyzed by a high-temperature catalytic oxidation method, and C_T was calculated from alkalinity–pH_T pairs and appropriate thermodynamic relationships. The results are used in a two-layer model of water mass exchange through the Strait, which includes the Atlantic inflow, the Mediterranean outflow and the interface layer in between. Our observations show a decrease of TOC and an increase of C_T concentrations from the surface to the bottom: 71–132 μM C and 2068–2150 μmol kg⁻¹ in the Surface Atlantic Water, 74–95 μM C and 2119–2148 μmol kg⁻¹ in the North Atlantic Central Water, 63–116 μM C and 2123–2312 μmol kg⁻¹ in the interface layer, and 61–78 μM C and 2307–2325 μmol kg⁻¹ in the Mediterranean waters. However, within the Mediterranean outflow, we found that the concentrations of carbon were higher at the western side of the Strait (75–78 μM C, 2068–2318 μmol kg⁻¹) than at the eastern side (61–69 μM C, 2082–2324 μmol kg⁻¹). This difference is due to the mixing between the Atlantic inflow and the Mediterranean outflow on the west of the Strait, which results in a flux of organic carbon from the inflow to the outflow and an opposite flux of inorganic carbon. We estimate that the TOC input from the Atlantic Ocean to the Mediterranean Sea through the Strait of Gibraltar varies from (0.97±0.8)10⁴ to (1.81±0.90)10⁴ mol C s⁻¹ (0.3×10¹² to 0.56×10¹² mol C yr⁻¹), while outflow of inorganic carbon ranges from (12.5±0.4)10⁴ to (15.6±0.4)10⁴ mol C s⁻¹ (3.99–4.90×10¹² mol C yr⁻¹). The high variability of carbon exchange within the Strait is due to the variability of vertical mixing between inflow and outflow along the Strait. The prevalence of organic carbon inflow and inorganic carbon outflow shows the Mediterranean Sea to be a basin of active remineralization of organic material.     

Mesozooplankton prey preference and grazing impact in the western Arctic Ocean

The role of mesozooplankton as consumers and transformers of primary and secondary production in the Beaufort and Chukchi Seas was examined during four cruises in spring and summer of both 2002 and 2004 as part of the western Arctic Shelf–Basin Interactions (SBI) program. Forty-seven grazing experiments using dominant mesozooplankton species and life stages were conducted at locations across the shelf, slope, and basin of the Chukchi and Beaufort Seas to measure feeding rates on both chlorophyll and microzooplankton and to determine mesozooplankton prey preferences.Mesozooplankton biomass was at all times dominated by life stages of four copepod taxa: Calanus glacialis, Calanus hyperboreus, Metridia longa, and Pseudocalanus spp. Significant interannual, seasonal, regional, between species and within species differences in grazing rates were observed. Overall, the dominant zooplankton exhibited typical feeding behavior in response to chlorophyll concentration that could be modeled using species and life-stage specific Ivlev functions. Microzooplankton were preferred prey at almost all times, with the strength of the preference positively related to the proportion of microzooplankton prey availability. Average mesozooplankton grazing impacts on both chlorophyll standing stock (0.6±0.5% d⁻¹ in spring, 5.1±6.3% d⁻¹ in summer) and primary production (12.8±11.8% d⁻¹ in spring, 27.6±24.5% d⁻¹ in summer) were quite low and varied between shelf, slope, and basin. Coincident microzooplankton grazing experiments [Sherr, E.B., Sherr, B.F., Hartz, A.J., 2009. Microzooplankton grazing impact in the Western Arctic Ocean. Deep-Sea Research II] were conducted at most stations. Together, microzooplankton–mesozooplankton grazing consumed only 44% of the total water-column primary production, leaving more than half directly available for local export to the benthos or for offshore transport into the adjacent basin.     

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.     

Structure and mechanisms of the Arabian Sea variability during the winter monsoon

In the southern Arabian Sea (between the Equator and 10°N), the shoaling of isotherms at subsurface levels (20 °C isotherm depth is located at ∼90 m) leads to cooling at 100 m by 2–3 °C relative to surrounding waters during the winter monsoon. The annual and interannual variations of this upwelling zone, which we call the Arabian Sea dome (ASD), are studied using results from an eddy-permitting ocean general circulation model in conjunction with hydrography and TOPEX/ERS altimeter data. The ASD first appears in the southeastern Arabian Sea during September–October, maturing during November–December to extend across the entire southern Arabian Sea (along ∼5°N). It begins to weaken in January and dissipates by March in the southwestern Arabian Sea. From the analysis of heat-budget balance terms and a pair of model control experiments, it is shown that the local Ekman upwelling induced by the positive wind-stress curl of the winter monsoon generates the ASD in the southeastern Arabian Sea. The ASD decays due to the weakening of the cyclonic curl of the wind and the westward penetration of warm water from the east (Southern Arabian Sea High). The interannual variation of the ASD is governed by variations in the Ekman upwelling induced by the cyclonic wind-stress curl. Associated with the unusual winds during 1994–1995 and 1997–1998 Indian Ocean dipole (IOD) periods, the ASD failed to develop. In the absence of the ASD during the IOD events, the 20 °C isotherm depth was 20–30 m deeper than normal in the southern Arabian Sea resulting in a temperature increase at 97 m of 4–5 °C. An implication is that the SST evolution in the southern Arabian Sea during the winter monsoon is primarily controlled by advective cooling: the shoaling of isotherms associated with the ASD leads to SST cooling.     

Distribution and burial of organic carbon in sediments from the Indian Ocean upwelling region off Java and Sumatra,Indonesia

Sediments were sampled and oxygen profiles of the water column were determined in the Indian Ocean off west and south Indonesia in order to obtain information on the production, transformation, and accumulation of organic matter (OM). The stable carbon isotope composition (δ¹³C_org) in combination with C/N ratios depicts the almost exclusively marine origin of sedimentary organic matter in the entire study area. Maximum concentrations of organic carbon (C_org) and nitrogen (N) of 3.0% and 0.31%, respectively, were observed in the northern Mentawai Basin and in the Savu and Lombok basins. Minimum δ¹⁵N values of 3.7‰ were measured in the northern Mentawai Basin, whereas they varied around 5.4‰ at stations outside this region. Minimum bottom water oxygen concentrations of 1.1 mL L^?1, corresponding to an oxygen saturation of 16.1%, indicate reduced ventilation of bottom water in the northern Mentawai Basin. This low bottom water oxygen reduces organic matter decomposition, which is demonstrated by the almost unaltered isotopic composition of nitrogen during early diagenesis. Maximum C_org accumulation rates (CARs) were measured in the Lombok (10.4 g C m^?2 yr^?1) and northern Mentawai basins (5.2 g C m^?2 yr^?1). Upwelling-induced high productivity is responsible for the high CAR off East Java, Lombok, and Savu Basins, while a better OM preservation caused by reduced ventilation contributes to the high CAR observed in the northern Mentawai Basin. The interplay between primary production, remineralisation, and organic carbon burial determines the regional heterogeneity. CAR in the Indian Ocean upwelling region off Indonesia is lower than in the Peru and Chile upwellings, but in the same order of magnitude as in the Arabian Sea, the Benguela, and Gulf of California upwellings, and corresponds to 0.1–7.1% of the global ocean carbon burial. This demonstrates the relevance of the Indian Ocean margin off Indonesia for the global OM burial.     

Isotopic and elemental indicators of nutrient sources and status of coastal habitats in the Caribbean Sea,Yucatan Peninsula,Mexico

Nutrient inputs associated with coastal population growth threaten the integrity of coastal ecosystems around the globe. In order to assess the threat posed by rapid growth in tourism, we analyzed the nutrient concentrations as well as the δ¹⁵N of NO₃⁻ and macrophytes to detect wastewater nitrogen (N) at 6 locations along a groundwater-dominated coastal seagrass bed on the Caribbean coast of Mexico. We predicted that locations with greater coastal development would have higher concentrations of dissolved inorganic nitrogen (DIN) and phosphorus (P), as well as δ¹⁵N of NO₃⁻, reflecting wastewater sources of N. However, concentrations of NO₃⁻ were not significantly different between developed (3.3 ± 5.3 μM NO₃⁻) and undeveloped (1.1 ± 0.7 μM) marine embayments. The most important control on DIN concentration appeared to be mixing of fresh and salt water, with DIN concentrations negatively correlated with salinity. The δ¹⁵N of NO₃⁻ was elevated at an inland pond (7.0 ± 0.42‰) and a hydrologically-connected tide pool (7.6 ± 0.57‰) approximately 1 km downstream of the pond. The elevated δ¹⁵N of NO₃⁻ at the pond was paralleled by high δ¹⁵N values of Cladophora sp., a ubiquitous green alga (10 ± 1‰). We hypothesize that inputs of nitrogen rich (NO₃⁻ > 30 μM) groundwater, characterized by ¹⁵N enriched signatures, flow through localized submarine groundwater discharges (SGD) and contribute to the elevated δ¹⁵N signatures observed in many benthic macrophytes. However, changes in nitrogen concentrations and isotope values over the salinity gradient suggest that other processes (e.g. denitrification) could also be contributing to the ¹⁵N enrichments observed in primary producers. More measurements are needed to determine the relative importance of nitrogen transformation processes as a source of ¹⁵N to groundwaters; however, it is clear that continued inputs of anthropogenic N via SGD have the potential to severely impact ecologically and economically valuable seagrass meadows and coral reefs along the Caribbean coast of Mexico.     

Bacterivory in the northwestern Indian Ocean during the intermonsoon – northeast monsoon period

Bacterial grazing loss rates were studied by radioactive labeling of natural bacteria with L-(4,5-³H) leucine and from the rate of disappearance of bacterial cells in the northwestern Indian Ocean. Bacterivory was measured in a mixed sample that had been combined from various depths across the euphotic zone. Experiments were performed on 26 occasions at 19 stations in the Gulf of Oman and the Arabian Sea during the intermonsoon–northeast monsoon period (November–December 1994). Combined uptake of radiolabeled bacteria (ULB) in 1–8 and 8–100 μm size fractions was somewhat lower than loss of label (LBL) measured in the bacterial fraction (0.2–1.0 μm), suggesting loss of radioactivity from the grazers due to metabolism. The less sensitive rate of disappearance of bacterial cells (LBC) was on average 51% higher than LBL estimates. Results from ULB and LBL measurements revealed that bacterivory was higher in the Gulf of Oman (average loss rate 4.1% h^-1) than in the Arabian Sea where rates were slightly higher inshore (1.7% h^-1) than in the central gyre. Heterotrophic nanoflagellates in the 1–8 μm size fractions were identified as the primary bacterivores. Microzooplankton (8–100 μm) accounted for 33% of total bacterivory in the Gulf of Oman but only 16% in the central Arabian Sea. Time-course experiments conducted at two stations indicated that diel changes in bacterivory may be substantial in the northwestern Indian Ocean.