The fate of production in the central Arctic Ocean - top-down regulation by zooplankton expatriates?

We estimated primary and bacterial production, mineral nutrients, suspended chlorophyll a (Chl), particulate organic carbon (POC) and nitrogen (PON), abundance of planktonic organisms, mesozooplankton fecal pellet production, and the vertical flux of organic particles of the central Arctic Ocean (Amundsen basin, 89-88° N) during a 3 week quasi-Lagrangian ice drift experiment at the peak of the productive season (August 2001). A visual estimate of ≈15% ice-free surface, plus numerous melt ponds on ice sheets, supported a planktonic particulate primary production of 50-150 mg C m⁻² d⁻¹ (mean 93 mg C m⁻² d⁻¹, n = 7), mostly confined to the upper 10 m of the nutrient replete water column. The surface mixed layer was separated from the rest of the water column by a strong halocline at 20 m depth. Phototrophic biomass was low, generally 0.03-0.3 mg Chl m⁻³ in the upper 20 m and <0.02 mg Chl m⁻³ below, dominated by various flagellates, dinoflagellates and diatoms. Bacterial abundance (typically 3.7-5.3 × 10⁵, mean 4.1 × 10⁵ cells ml⁻¹ in the upper 20 m and 1.3-3.7 × 10⁵, mean 1.9 × 10⁵ cells ml⁻¹ below) and Chl concentrations were closely correlated (r = 0.75). Mineral nutrients (3 μmol NO₃ l⁻¹, 0.45 μmol PO₄ l⁻¹, 4-5 μmol SiO₄ l⁻¹) were probably not limiting the primary production in the upper layer. Suspended POC concentration was ∼30-105 (mean 53) mg C m⁻³ and PON ∼5.4-14.9 (mean 8.2) mg N m⁻³ with no clear vertical trend. The vertical flux of POC in the upper 30-100 m water column was ∼37-92 (mean 55) mg C m⁻² d⁻¹ without clear decrease with depth, and was quite similar at the six investigated stations. The mesozooplankton biomass (≈2 g DW m⁻², mostly in the upper 50 m water column) was dominated by adult females of the large calanoid copepods Calanus hyperboreus and Calanus glacialis (≈1.6 g DW m⁻²). The grazing of these copepods (estimated via fecal pellet production rates) was ≈15 mg C m⁻² d⁻¹, being on the order of 3% and 20% of the expected food-saturated ingestion rates of C. hyperboreus and C. glacialis, respectively. The stage structure of these copepods, dominated by adult females, and their unsatisfied grazing capacity during peak productive period suggest allochthonous origin of these species from productive shelf areas, supported by their long life span and the prevailing surface currents in the Arctic Ocean. We propose that the grazing capacity of the expatriated mesozooplankton population would match the potential seasonal increase of primary production in the future decreased ice perspective, diminishing the likelihood of algal blooms.     

Contrasting oceanographic conditions and phytoplankton communities on the east and west coasts of Australia

The composition and dynamics of the phytoplankton communities and hydrographic factors that control them are described for eastern and western Australia with a focus on the Eastern Australian Current (EAC) and Leeuwin Current (LC) between 27.5° and 34.5°S latitude. A total of 1685 samples collected from 1996 to 2010 and analysed for pigments by high performance liquid chromatography (HPLC) showed the average TChla (monovinyl+divinyl chlorophyll a) concentration on the west coast to be 0.28±0.16 ??g L⁻¹ while it was 0.58±1.4 ??g L⁻¹ on the east coast. Both coasts showed significant decreases in the proportions of picoplankton and relatively more nanoplankton and microplankton with increasing latitude. On both coasts the phytoplankton biomass (by SeaWiFS) increased with the onset of winter. At higher latitudes (>27.5°S) the southeast coast developed a spring bloom (September) when the mean monthly, surface chlorophyll a (chla) concentration (by SeaWiFS) was 48% greater than on the south west coast. In this southern region (27.5-34.5°S) Synechococcus was the dominant taxon with 60% of the total biomass in the southeast (SE) and 43% in the southwest (SW). Both the SE and SW regions had similar proportions of haptophytes; ∼14% of the phytoplankton community. The SW coast had relatively more pelagophytes, prasinophytes, cryptophytes, chlorophytes and less bacillariophytes and dinophytes. These differences in phytoplankton biomass and community composition reflect the differences in seasonality of the 2 major boundary currents, the influence this has on the vertical stability of the water column and the average availability of nutrients in the euphotic zone. Seasonal variation in mixed layer depth and upwelling on the west coast appears to be suppressed by the Leeuwin Current. The long-term depth averaged (0-100 m) nitrate concentration on the west coast was only 14% of the average concentration on the east coast. Redfield ratios for NO₃:SiO₂:PO₄ were 6.5:11.9:1 on the east coast and 2.2:16.2:1 on the west coast. Thus new production (nitrate based) on the west coast was likely to be substantially more limited than on the eastcoast. Short term (hourly) rates of vertical mixing were greater on the east coast. The more stable water column on the west coast produced deeper subsurface chlorophyll a maxima with a 25% greater proportion of picoeukaryotes.     

Dynamics of heterotrophic dinoflagellates off the Pearl River Estuary,northern South China Sea

Variations in abundance, biomass, vertical profile and cell size of heterotrophic dinoflagellates (HDFs) between summer and winter and its controlling factors were studied in the northern South China Sea (SCS). It was found that HDF abundance and carbon biomass were 4–102 × 10³ cells L⁻¹ and 0.34–12.3 mg C L⁻¹ in winter (February 2004), respectively, while they were 2–142 × 10³ cells L⁻¹ and 0.22–31.4 μg C L⁻¹ in summer (July, 2004), respectively, in the northern SCS. HDF abundance and carbon biomass decreased from the estuary to inshore and then offshore. Vertical profiles of HDF abundance were heterogeneous, which accorded well with that of chlorophyll a (Chl.a). Higher abundance of HDFs was often observed at a depth of 30–70 m offshore waters, matching well with the Chl.a maximum, while it showed high abundance at the surface in some coastal and estuary stations. Small HDFs (≤20 μm) dominated the assemblage in term of abundance accounting for more than 90%. However, large HDFs (>20 μm) generally contributed equally in terms of carbon biomass, accounting for 47% on average. HDFs showed different variation patterns for the different study regions; in the estuarine and continental shelf regions, abundance and biomass values were higher in summer than those in winter, while it was the reverse pattern for the slope waters. Hydrological factors (e.g. water mass, river outflow, monsoon and eddies) associated with biological factors, especially the size-fractionated Chl.a, seemed to play an important role in regulating HDF distribution and variations in the northern South China Sea.     

The planktonic food web of the Bizerte lagoon (south-western Mediterranean) during summer: I. Spatial distribution under different anthropogenic pressures

The structure and the trophic interactions of the planktonic food web were investigated during summer 2004 in a coastal lagoon of south-western Mediterranean Sea. Biomasses of planktonic components as well as bacterial and phytoplankton production and grazing by microzooplankton were quantified at four stations (MA, MB, MJ and R) inside the lagoon. Station MA was impacted by urban discharge, station MB was influenced by industrial activity, station MJ was located in a shellfish farming sector, while station R represented the lagoon central area. Biomasses and production rates of bacteria (7–33 mg C m⁻³; 17.5–35 mg C m⁻³ d⁻¹) and phytoplankton (80–299 mg C m⁻³; 34–210 mg C m⁻³ d⁻¹) showed high values at station MJ, where substantial concentrations of nutrients (NO₃⁻ and Si(OH)₄) were found. Microphytoplankton, which dominated the total algal biomass and production (>82%), were characterized by the proliferation of several chain-forming diatoms. Microzooplankton was mainly composed of dinoflagellates (Torodinium, Protoperidinium and Dinophysis) and aloricate (Lohmaniellea and Strombidium) and tintinnid (Tintinnopsis, Tintinnus, Favella and Eutintinnus) ciliates. Higher biomass of these protozoa (359 mg C m⁻³) was observed at station MB, where large tintinnids were encountered. Mesozooplankton mainly represented by Calanoida (Acartia, Temora, Calanus, Eucalanus, Paracalanus and Centropages) and Cyclopoida (Oithona) copepods, exhibited higher and lower biomasses at stations MA/MJ and MB, respectively. Bacterivory represented only 35% of bacterial production at stations MB and R, but higher fractions (65–70%) were observed at stations MA and MJ. Small heterotrophic flagellates and aloricate ciliates seemed to be the main controllers of bacteria. Pico- and nanophytoplankton represented a significant alternative carbon pool for micrograzers, which grazing represented 67–90% of pico- and nano-algal production in all stations. Microzooplankton has, however, a relatively low impact on microphytoplankton, as ≤45% of microalgal production was consumed in all stations. This implies that an important fraction of diatom production would be channelled by herbivorous meso-grazers to higher consumers at stations MA and MJ where copepods were numerous. Most of the microalgal production would, however, sink particularly at station MB where copepods were scare. These different trophic interactions suggest different food web structures between stations. A multivorous food web seemed to prevail in stations MJ and MA, whereas microbial web was dominant in the other stations.     

Influence of river discharge on plankton metabolic rates in the tropical monsoon driven Godavari estuary,India

To examine the influence of river discharge on plankton metabolic balance in a monsoon driven tropical estuary, daily variations in physico-chemical and nutrients characteristics were studied over a period of 15 months (September 2007 to November 2008) at a fixed location (Yanam) in the Godavari estuary, India. River discharge was at its peak during July to September with a sharp decrease in the middle of December and complete cessation thereafter. Significant amount of dissolved inorganic nitrogen (DIN, of 22–26 μmol l⁻¹) and dissolved inorganic phosphate (DIP, of 3–4 μmol l⁻¹) along with suspended materials (0.2–0.5 g l⁻¹) were found at the study region during the peak discharge period. A net heterotrophy with low gross primary production (GPP) occurred during the peak discharge period. The Chlorophyll a (Chl a) varied between 4 and 18 mg m⁻³ that reached maximum levels when river discharge and suspended loads decreased by >75% compared to that during peak period. High productivity was sustained for about one and half months during October to November when net community production (NCP) turned from net heterotrophy to autotrophy in the photic zone. Rapid decrease in nutrients (DIN and DIP by ∼15 and 1.4 μmol l⁻¹, respectively) was observed during the peak Chl a period of two weeks. Chl a in the post monsoon (October–November) was negatively related to river discharge. Another peak in Chl a in January to February was associated with higher nutrient concentrations and high DIN:DIP ratios suggest possible external supply of nitrogen into the system. The mean photic zone productivity to respiration ratio (P:R) was 2.38 ± 0.24 for the entire study period (September 2007–November 2008). Nevertheless, the ratio of GPP to the entire water column respiration was only 0.14 ± 0.02 revealing that primary production was not enough to support water column heterotrophic activity. The excess carbon demand by the heterotrophs could be met from the allochthonous inputs of mainly terrestrial origin. Assuming that the entire phytoplankton produced organic material was utilized, the additional terrestrial organic carbon supported the total bacterial activity (97–99%) during peak discharge period and 40–75% during dry period. Therefore, large amount of terrestrial organic carbon is getting decomposed in the Godavari estuarine system.     

Growth and grazing rate dynamics of major phytoplankton groups in an oligotrophic coastal site

There has been more attention to phytoplankton dynamics in nutrient-rich waters than in oligotrophic ones thus requiring the need to study the dynamics and responses in oligotrophic waters. Accordingly, phytoplankton community in Blanes Bay was overall dominated by Prymnesiophyceae, remarkably constant throughout the year (31 ± 13% Total chlorophyll a, Tchl a) and Bacillariophyta with a more episodic appearance (20 ± 23% Tchl a). Prasinophyceae and Synechococcus contribution became substantial in winter (Prasinophyceae = 30% Tchl a) and summer (Synechococcus = 35% Tchl a). Phytoplankton growth and grazing mortality rates for major groups were estimated by dilution experiments in combination with high pressure liquid chromatography and flow cytometry carried out monthly over two years. Growth rates of total phytoplankton (range = 0.30–1.91 d⁻¹) were significantly higher in spring and summer (μ > 1.3 d⁻¹) than in autumn and winter (μ ∼ 0.65 d⁻¹) and showed a weak dependence on temperature but a significant positive correlation with day length. Microzooplankton grazing (range = 0.03–1.4 d⁻¹) was closely coupled to phytoplankton growth. Grazing represented the main process for loss of phytoplankton, removing 60 ± 34% (±SD) of daily primary production and 70 ± 48% of Tchl a stock. Chla synthesis was highest during the Bacillarophyceae-dominated spring bloom (Chl a_synt = 2.3 ± 1.6 μg Chl a L⁻¹ d⁻¹) and lowest during the following post-bloom conditions dominated by Prymnesiophyceae (Chl a_synt = 0.23 ± 0.08 μg Chl a L⁻¹ d⁻¹). This variability was smoothed when expressed in carbon equivalents mainly due to the opposite dynamics of C:chl a (range = 11–135) and chl a concentration (range = 0.07–2.0 μg chl a L⁻¹). Bacillariophyta and Synechococcus contribution to C fluxes was higher than to biomass because of their fast-growth rate. The opposite was true for Prymnesiophyceae.     

Grazing impact of the copepod community in the Oyashio region of the western subarctic Pacific Ocean

The role of copepod grazing on the ecosystem dynamics in the Oyashio region, western subarctic Pacific was investigated during six cruises from June 2001 to June 2002. In situ grazing rates of the copepod community (CGR) were measured by the gut fluorescence method in respect to developmental stages of dominant species. In terms of biomass, more than 80% of the copepod community was dominated by six large calanoid species (Neocalanus cristatus, Neocalanus flemingeri, Neocalanus plumchrus, Eucalanus bungii, Metridia pacifica and Metridia okhotensis) throughout the year. Resulting from the observed pattern of the interzonal migrating copepods, the CGR in the Oyashio region was divided into three phases, i.e. spring (bloom), summer (post-bloom) and autumn-winter phase. During the spring bloom, late copepodites of the interzonal migrating species, N. cristatus, N. flemingeri and E. bungii appeared in the surface layer (0-50 m) to consume the production of the bloom, resulting in a high grazing rate of the copepod community (7.9 mg Chl m⁻² d⁻¹), though its impact on phytoplankton community was low due to the high primary productivity. During the post-bloom period, although the copepod community which was dominated by N. cristatus, N. plumchrus, M. pacifica and newly recruited E. bungii still maintained a high biomass, the CGR was generally lower (1.8-2.6 mg Chl m⁻² d⁻¹ for June and August 2001), probably due to the lower availability of phytoplankton. Nevertheless, the highest CGR was also observed during this period (10.5 mg Chl m⁻² d⁻¹ in June 2002). The high CGR on autotrophic carbon accounted for 69% of the primary production, suggesting that the copepod community in the Oyashio region potentially terminates the phytoplankton bloom. Abundant occurrence of young E. bungii, which is a characteristic phenomenon in the Oyashio region, was largely responsible for the high grazing pressure in June 2002 suggesting that success of reproduction, growth, and survival in E. bungii during the spring bloom is an important factor in controlling phytoplankton abundance during the post-bloom season. During autumn and winter, CGR was the lowest in the year (0.29-0.38 mg Chl. m⁻² d⁻¹) due to the disappearance of the interzonal migrating copepods from the surface layer. Diel migrant M. pacifica was the most important grazer during this period. The annual ingestion of the copepod community is estimated as 37.7 gC m⁻² on autotrophic carbon (converted using C:Chl ratio of 30) or 137.9 gC m⁻² on suspended particles (using C:Chl ratio of in situ value, 58-191), accounting for 13% and 46% of annual primary production, respectively. This study confirms that copepod grazing is an important pathway in carbon flow in the Oyashio region and in particular their role in the phytoplankton dynamics is significant for the termination of the spring bloom.     

Microbial plankton abundance and heterotrophic activity across the Central Atlantic Ocean

The role of microorganisms in the transfer of carbon of marine systems is very important in open oligotrophic oceans. Here, we analyze the picoplankton structure, the heterotrophic bacterioplankton activity, and the predator-prey relationships between heterotrophic bacteria and nanoflagellates during two large scale cruises in the Central Atlantic Ocean (∼29°N to ∼40°S). Latitud cruises were performed in 1995 between March-April and October-November. During both cruises we crossed the regions of different trophic statuses; where we measured different biological variables both at the surface and at the deep chlorophyll maximum (DCM). The concentration of chlorophyll a varied between 0.1 and 0.8 mg m⁻³, the abundance of heterotrophic bacteria varied between <1.0 × 10⁵ and >1.0 × 10⁶ cells ml⁻¹, and that of heterotrophic nanoflagellates between <100 and >1.0 × 10⁴ cells ml⁻¹. The production of heterotrophic bacteria varied more than three orders of magnitude between <0.01 and 24 μgC L⁻¹ d⁻¹; and the growth rates were in the range <0.01-2.1 d⁻¹. In the Latitud-II cruise, Prochlorococcus ranged between <10³ and >3 × 10⁵ cells ml⁻¹, Synechococcus between <100 and >1.0 × 10⁴ cells ml⁻¹, and picoeukaryotes between <100 and >10⁴ cells ml⁻¹.Two empirical models were used to learn more about the relationship between heterotrophic bacteria and nanoflagellates. Most bacterial production was ingested when this production was low, the heterotrophic nanoflagellates could be controlled by preys during Latitud-I cruise at the DCM, and by predators in the surface and in the Latitud-II cruise. Our results were placed in context with others about the structure and function of auto- and heterotrophic picoplankton and heterotrophic nanoplankton in the Central Atlantic Ocean.     

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.     

Dissolved organic carbon in the southern Baltic Sea: Quantification of factors affecting its distribution

During a cruise of r/v ‘Oceania’ in May 2006, seven vertical dissolved organic carbon (DOC) concentration profiles were produced against a background of CTD, chlorophyll a (chl a) and phaeopigment concentration profiles. The results indicate distinct vertical and spatial DOC fluctuations, ranging from 248 ± 7 μmol C dm⁻³ at 70 m depth at the westernmost station G/06 to 398 ± 5 μmol C dm⁻³ at 5 m depth at station A/06 in the western Gulf of Gdańsk. DOC concentrations were the highest at 10 m depth, where phytoplankton activity was relatively intensive, as reflected by the active chl a concentration distribution. DOC concentrations decreased towards the sea bottom.     

Spatial and temporal variability of microplankton and detritus, and their export to the shelf sediments in the upwelling area off Concepción, Chile (∼36°S), during 2002-2005

Seasonal variations in diversity and biomass of diatoms, tintinnids, and dinoflagellates and the contribution of microplankton and faecal material to the vertical flux of particulates were investigated at one time series station T (station 18) between 2002 and 2005 and at a grid of stations during November 2004 in the coastal and oceanic area off Concepción (36°S), Chile. The variations were analysed in relation to water column temperature, dissolved oxygen, nutrient concentration, offshore Ekman transport, and chlorophyll-a concentration. Abundance was estimated as cell numbers per litre and biomass in terms of biovolume and carbon units.A sharp decrease with depth was observed in the abundance of both phytoplankton and microzooplankton during the whole annual cycle; over 70% of their abundance was concentrated in the upper 10 m of the water column. Also, a clear seasonality in microplankton distribution was observed at station T, with maxima for diatoms, tintinnids, and dinoflagellates every summer (centred on January) from 2002 to 2005.On the grid of stations, the maximum integrated (0-50 m) micro-phytoplankton abundances (>1 × 10⁹ cells m⁻²) occurred at the coastal stations, an area directly influenced by upwelling. A similar spatial distribution was observed for the integrated (0-200 m) faecal carbon (with values up to 632 mg C m⁻²). Tintinnids were distributed in all the first 300 miles from the coast and dinoflagellates were more abundant in oceanic waters.At station T, the average POC export production (below 50 m depth) was 16.6% (SD = 17%; range 2-67%; n = 16). The biological-mediated fluxes of carbon between the upper productive layer and the sediments of the continental shelf off Concepción depend upon key groups of phytoplankton (Thalassiosira spp., Chaetoceros spp.) and zooplankton (euphausiids) through the export of either cells or faecal material, respectively.     

Nanoplanktonic assemblages in the upwelling area off Concepción (∼36°S), central Chile: Abundance, biomass, and grazing potential during the annual cycle

The structure and functioning of nanoplanktonic assemblages in coastal upwelling areas have usually been overlooked in explorations of the productivity of these areas. As part of a multidisciplinary, time-series station in the coastal area off Concepción, seasonal variations (upwelling and non-upwelling) in the abundance and biomass of these assemblages were investigated. Hydrographic measurements and biological samples were taken monthly over a 2-year period (18 August 2004-28 July 2006). Nanoflagellates dominated the total integrated abundance (3-317 × 10⁹ cells m⁻²; 0-80 m). Diatoms and dinoflagellates usually contributed to a lesser degree (<20%) but sporadically made important contributions to the total integrated nanoplankton biomass (0.02-10.6 g C m⁻²). Most of the nanoplankton was concentrated in surface waters (<30 m) during all the samplings and no seasonal differences in abundance or biomass were found in this layer, although the mean values and dispersions around them were highest during the upwelling period along with maximum integrated (0-80 m) chlorophyll-a values, as total or in the <20 μm fraction. Changes in nanoplankton abundance were significantly but weakly (r < 0.4) correlated with changes in the hydrographic variables; the highest correlation values were positive for temperature and oxygen, factors that varied with depth and date. The potential grazing rates of heterotrophic nano-predators (flagellates and dinoflagellates) on prokaryotic prey, estimated with a generic model, ranged from 3 to 242 bacterioplankton predator⁻¹ h⁻¹ and from 0.1 to 14 cyanobacteria predator⁻¹ h⁻¹. Our results imply a small impact of seasonal hydrographic variability on the abundance and biomass of nanoplanktonic assemblages and suggest that grazing by nanoheterotrophs might control the prokaryotic picoplankton populations in the upwelling area off Concepción.     

The annual silica cycle of the North Pacific subtropical gyre

Silica cycling in the upper 175 m of the North Pacific Subtropical Gyre was examined over a two year period (January 2008-December 2009) at the Hawaii Ocean Time-series (HOT) station ALOHA. Silicic acid concentrations in surface waters ranged from 0.6 to 1.6 ??M, exhibiting no clear seasonal trends. Biogenic silica concentrations and silica production rates increased by an order of magnitude each summer following stratification of the upper 50 m reaching values of 157 nmol Si L⁻¹ and 81 nmol Si L⁻¹ d⁻¹, in 2008 and 2009, respectively. Sea surface height anomalies together with analyses of variability in isothermal surfaces at 150-175 m indicated that the summer periods of elevated biogenic silica were associated with anticyclonic mesoscale features during both years. Lithogenic silica concentrations increased in the spring during the known period of maximum atmospheric dust concentrations with maximum values of 36 nmol Si L⁻¹ in the upper 10 m. Dust deposition would enhance levels of dissolved iron in surface waters, but there was no response of diatom biomass or silica production to increases in near-surface ocean lithogenic silica concentrations suggesting iron sufficiency of diatom silica production rates.Low ambient silicic acid concentrations restricted silica production rates to an average of 43% of maximum potential rates. Si sufficiency only occurred during the summer period when diatom biomass was elevated suggesting that bloom diatoms are adapted to exploit low silicic acid concentrations. Annual silica production at HOT is estimated to be 63 mmol Si m⁻² a⁻¹ with summer blooms contributing 29% of the annual total. Diatoms are estimated to account for 3-7% of total phytoplankton primary productivity, but 9-20% of organic carbon export confirming past suggestions that diatoms are relatively minor contributors to primary productivity and autotrophic biomass, but important contributors to new and export production in oligotrophic open-ocean ecosystems.Annual silica production at HOT is nearly 4-fold lower than estimates at the Bermuda Atlantic Time-series Study (BATS) site in the Sargasso Sea from the 1990s, but annual silica export at the base of the euphotic zone is similar between the two gyres indicating very different balances between silica production and its loss in surface waters. On a relative basis, BATS is a more productive system with respect to silica, where biogenic silica is recycled with high efficiency in surface waters; in contrast the NPSG is a lower productivity system with respect to silica, but where lower recycling efficiency leads to a much higher fraction of new silica production. The two gyres show contrasting long-term trends in diatom biomass as biogenic silica concentrations at HOT have been increasing since 1997, but they have been decreasing at BATS suggesting very different forcing of decadal trends in the contribution of diatoms in carbon cycling between these gyres. Combining the data from both gyres indicates that globally subtropical gyres produce 13 Tmol Si a⁻¹, which is only 51% of previous estimates reducing the contribution of subtropical gyres to 5-7% of global annual marine silica production.     

Assessment of carbon- and fluorescence-based primary productivity in Ariake Bay,southwestern Japan

A comparative account of primary productivity (PP), in the characteristically turbid and highly dynamic waters of Ariake Bay, measured by ¹³C uptake and fast repetition rate fluorometer (FRRF) was conducted to ensure compatibility between the two methods. Estimates from both methods depicted strong linearity for both short-term (r² > 0.90) and daily (r² = 0.42–0.93) measurements, except in the near-surface (∼0 m) layer. ¹³C-based short-term (1 h; in situ) PP estimates showed similar magnitudes and trend with the instantaneous PP measured by FRRF concurrently. Whereas, unlike short-term measurements, the daily PP estimates from both methods showed large difference, with FRRF-based time integrated daily PP resulting in 1.09–1.82 times higher than the carbon-based daily (24 h; simulated in situ) PP. This difference between daily PP estimates was mainly due to: (1) the temporal variation of water column chlorophyll a (Chl a) because of frequent moving of water mass, and (2) the dissimilarity in ambient light field conditions between the two methods. Results revealed that considering the above two environmental factors invariable over a daylength, fairly close approximation of daily PP, compared to ¹³C-based daily PP, could be obtained from FRRF. Hence, FRRF-based daily PP can be considered as more realistic in this highly dynamic water body like Ariake Bay where water column parameters are subjected to strong temporal variation. The relationship between Chl a-specific photosynthetic rate (P^B) and the corresponding photosynthetically active radiation (PAR) in the water column (PAR–P^B relationship) was found to be linear for FRRF and curvilinear for ¹³C-based measurements in the near-surface layer, for the same intensities of incident PAR, and this is thought to be the primary basis for the higher difference in PP estimates at the near-surface layer. Considering the minor variations in FRRF-based time series of PAR–P^B relationships, a combined and/or instantaneous PAR–P^B relationship in combination with incubation Chl a and light field condition was used to obtain fairly close estimates of daily water column integrated PP from FRRF.     

Dynamics and sources of suspended particulate organic matter in the Marennes-Oléron oyster farming bay: Insights from stable isotopes and microalgae ecology

The aim of this study was to distinguish between sources of the complex variety of Marennes-Oléron Bay suspended particulate organic matter (SPOM) contributing to the tropho-dynamics of the Marennes-Oléron oyster farming bay. Basic biomarkers (Chl a, C/N and POC/Chl a ratios), carbon and nitrogen stable isotopes from SPOM were analyzed and the microalgae community was characterized. The sampling strategy was bimonthly from March 2002 to December 2003; samples were taken from an intertidal mudflat. Four main sources contributed to the SPOM pool: terrigenous input from rivers, neritic phytoplankton, resuspended microphytobenthos and periodic inputs from intertidal Zostera noltii meadows. Seasonal fluctuations were observed in both years of the study period: (1) SPOM collected in the spring of 2002 (δ¹³C = −25‰ to −23‰) was mainly composed of fresh estuarine inputs; (2) SPOM from the summer and fall of 2002 and 2003 was predominantly neritic phytoplankton (δ¹³C = − 22‰ to −19‰); (3) SPOM from the winter of 2002, spring of 2003 and winter of 2003 (δ¹³C = −21 to −23‰) was composed of a mixture of decayed terrigenous river inputs and pelagic phytoplankton, which was predominantly resuspended microphytobenthos. In the summer of 2003—the warmest summer on record in southern France and Europe—SPOM was particularly enriched for ¹³C, with δ¹³C values ranging from −14‰ to −12‰. Pulses in δ¹³C values, indicative of ¹³C-enriched decaying materials, extended into the fall. These were attributed to benthic intertidal inputs, including both resuspended microphytobenthos and Z. noltii detritus. Changes in SPOM sources in Marennes-Oléron Bay may lead to differences in the quality of the trophic environment available for reared oysters.     

Pteropods in Southern Ocean ecosystems

To date, little research has been carried out on pelagic gastropod molluscs (pteropods) in Southern Ocean ecosystems. However, recent predictions are that, due to acidification resulting from a business as usual approach to CO₂ emissions (IS92a), Southern Ocean surface waters may begin to become uninhabitable for aragonite shelled thecosome pteropods by 2050. To gain insight into the potential impact that this would have on Southern Ocean ecosystems, we have here synthesized available data on pteropod distributions and densities, assessed current knowledge of pteropod ecology, and highlighted knowledge gaps and directions for future research on this zooplankton group.Six species of pteropod are typical of the Southern Ocean south of the Sub-Tropical Convergence, including the four Thecosomes Limacina helicina antarctica, Limacina retroversa australis, Clio pyramidata, and Clio piatkowskii, and two Gymnosomes Clione limacina antarctica and Spongiobranchaea australis. Limacina retroversa australis dominated pteropod densities north of the Polar Front (PF), averaging 60 ind m⁻³ (max = 800 ind m⁻³) and 11% of total zooplankton at the Prince Edward Islands. South of the PF L. helicina antarctica predominated, averaging 165 ind m⁻³ (max = 2681 ind m⁻³) and up to >35% of total zooplankton at South Georgia, and up to 1397 ind m⁻³ and 63% of total zooplankton in the Ross Sea. Combined pteropods contributed <5% to total zooplankton in the Lazarev Sea, but 15% (max = 93%) to macrozooplankton in the East Antarctic. In addition to regional density distributions we have synthesized data on vertical distributions, seasonal cycles, and inter-annual density variation.Trophically, gymnosome are specialist predators on thecosomes, while thecosomes are considered predominantly herbivorous, capturing food with a mucous web. The ingestion rates of L. retroversa australis are in the upper range for sub-Antarctic mesozooplankton (31.2-4196.9 ng pig ind⁻¹ d⁻¹), while those of L. helicina antarctica and C. pyramidata are in the upper range for all Southern Ocean zooplankton, in the latter species reaching 27,757 ng pig ind⁻¹ d⁻¹ and >40% of community grazing impact. Further research is required to quantify diet selectivity, the effect of phytoplankton composition on growth and reproductive success, and the role of carnivory in thecosomes.Life histories are a significant knowledge gap for Southern Ocean pteropods, a single study having been completed for L. retroversa australis, making population studies a priority for this group. Pteropods appear to be important in biogeochemical cycling, thecosome shells contributing >50% to carbonate flux in the deep ocean south of the PF. Pteropods may also contribute significantly to organic carbon flux through the production of fast sinking faecal pellets and mucous flocs, and rapid sinking of dead animals ballasted by their aragonite shells. Quantification of these contributions requires data on mucous web production rates, egestion rates, assimilation efficiencies, metabolic rates, and faecal pellet morphology for application to sediment trap studies.Based on the available data, pteropods are regionally significant components of the Southern Ocean pelagic ecosystem. However, there is an urgent need for focused research on this group in order to quantify how a decline in pteropod densities may impact on Southern Ocean ecosystems.     

Close coupling between ammonium uptake by phytoplankton and excretion by Antarctic krill, Euphausia superba

In this study we examined the hypothesis that, under conditions of replete macronutrients and iron in the Southern Ocean, phytoplankton abundance and specific N uptake rates are influenced strongly by the processes of grazing and NH₄ regeneration. NH₄ and NO₃ uptake rates by marine phytoplankton were measured to the northeast and northwest of the island of South Georgia during January-February 1998. Mean specific uptake rate for NO₃ (vNO₃) was 0.0026 h⁻¹ (range 0.0013-0.0065 h⁻¹) and for NH₄ (vNH₄) was 0.0097 h⁻¹ (0.0014-0.0376 h⁻¹). vNH₄ was related positively with NH₄ availability, which ranged from 0.1 to 1.5 mmol m⁻³ within the upper mixed layer. Ambient NH₄ concentrations and vNH₄ were both positively related to local krill biomass values, computed from mean values along acoustic transect segments within 2 km of the uptake measurement stations. These biomass values ranged from ∼1 g krill fresh mass m⁻² in the northwest to >4 kg krill wet mass m⁻² in the northeast. In contrast to the variability found with NH₄ concentrations and uptake rates, vNO₃ was more uniform across the sampling sites. Under these conditions, increasing NH₄ concentration appeared to represent an additional N resource. However, high vNH₄ tended to be found for stations with lower phytoplankton standing stocks, across a total range of 0.24-20 mg chlorophyll a m⁻³. These patterns suggest a coupling between phytoplankton biomass, vNH₄ and krill in this region of variable but high krill biomass. Locally high concentrations of krill in parts of the study area appeared to have two opposing effects. On the one hand they could graze down phytoplankton stocks, but on the other hand, their NH₄ excretion supported enhanced uptake rates by the remaining, ungrazed cells.     

Particulate suspended matter concentrations in the Bahía Blanca Estuary,Argentina: Implication for the development of phytoplankton blooms

The inner zone of the Bahía Blanca Estuary is shallow, nutrient-rich and turbid. Tidal energy and water turbulence strongly affect the water column resulting in a well-mixed structure and high concentrations of suspended sediment. The phytoplankton community is mostly dominated by diatoms and the annual pattern has been characterized by a recurrent winter-early spring bloom. Here, we investigated to what extent the temporal variations of suspended particulate matter (SPM) regulate the phytoplankton blooms in the head of the estuary by light-limitation. Sampling was done on a fortnightly basis (weekly during the blooming season) at a fixed station in the inner zone of the estuary from January 2007 to February 2008. SPM concentrations and light extinction coefficients (k) in the water column were significantly correlated and showed relatively lower values during the phytoplankton maximal biomass levels. During winter, SPM and k reached values of 23.6 mg l⁻¹ and 0.17 m⁻¹ which were significantly lower than the annual means of 77.6 mg l⁻¹ and 2.94 m⁻¹, respectively. The particulate organic matter (POM) concentration was significantly correlated with the calculated phytoplankton biomass although the contribution of the latter to the total POM was rather low. Both, POM and biomass, had maximal values during winter (21.8 mg l⁻¹ and 393.5 μg C l⁻¹) and mid summer (24.3 mg l⁻¹ and 407.0 μg C l⁻¹), with cell densities up to 8 × 10⁶ cells l⁻¹ and chlorophyll a up to 24.6 μg l⁻¹. Our results suggest that the decrease of SPM concentrations in the water column with a concomitant increase in the penetration of solar radiation seems to be one of the main causes for the development of the phytoplankton winter bloom in the Bahía Blanca Estuary.     

Spatial and temporal variability in coccolithophore abundance and production of PIC and POC in the NE subarctic Pacific during El Niño (1998), La Niña (1999) and 2000

Seasonal variations in coccolithophore abundance, chlorophyll, nutrients and production of particulate organic and inorganic carbon (POC and PIC) were determined along a coastal to oceanic east-west transect (Line P) culminating at Ocean Station Papa in the northeastern subarctic Pacific between 1998 and 2000. Offshore stations generally exhibited low seasonality in chlorophyll concentrations, with moderate seasonality in POC production. Near shelf stations showed a similar pattern to offshore stations, but were also characterized by sporadic events of higher POC productivity. During the 1998 El Niño, June was characterized by low chlorophyll and POC productivity along the transect, presumably as a result of depleted surface nitrate. In contrast, during the 1999 La Niña, and in 2000, higher POC productivity and surface nitrate occurred along the transect in June. Chlorophyll and POC productivity were similar in late summer in all 3 years. The coccolithophore population was usually numerically dominated by Emiliania huxleyi, particularly in June. Along the transect, abundance of coccolithophores was much higher in June during the 1998 El Niño (mean of 221 cells ml⁻¹) than in the 1999 La Niña (mean of 40 cells ml⁻¹), with their abundance in late summers of both years being very low. Abundances were even higher along the transect in June and the late summer of 2000 with sporadic ‘blooms’ of >1000 cells ml⁻¹ at some stations (cruise averages 395 and 552 cell ml⁻¹, respectively). Production rates of PIC did not consistently correlate with areas of high coccolithophore abundance. PIC production was high (100-250 mg C m⁻² d⁻¹) along the transect during June 1998, and low (1-40 mg C m⁻² d⁻¹) during both winters, June 1999 and during late summers of 1998 and 1999. The year 2000 was more complicated, with high rates of PIC production accompanying high abundance of coccolithophores in late summer, but lower rates of PIC production accompanying high coccolithophore numbers in June. Our data suggest that the abundance of coccolithophores and the production rates of PIC in the subarctic are higher than previously thought. Occasional PIC:POC production ratios of 1 or greater in 1998 and 2000 suggest that coccolithophores in this region could have a significant impact on the efficiency of the biological carbon pump.     

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.