Distribution of bacterial abundance and cell-specific nucleic acid content in the Northeast Pacific Ocean

We tested the idea that bacterial cells with high nucleic acid content (HNA cells) are the active component of marine bacterioplankton assemblages, while bacteria with low nucleic acid content (LNA cells) are inactive, with a large data set (>1700 discrete samples) based on flow cytometric analysis of bacterioplankton in the Northeast Pacific Ocean off the coast of Oregon and northern California, USA. Samples were collected in the upper 150 m of the water column from the coast to 250 km offshore during 14 cruises from March 2001 to September 2003. During this period, a wide range of trophic states was encountered, from dense diatom blooms (chlorophyll-a concentrations up to 43 μg l⁻¹) at shelf stations during upwelling season (March–September) to lower chlorophyll-a concentrations (0.1–5 μg l⁻¹) during winter (November–February) and at basin stations (>1700 m depth). We found only weakly positive relations of log total bacterial abundance to log chlorophyll-a concentration (as a proxy for availability of organic substrate), and of HNA bacteria as a fraction of total bacteria to log chlorophyll-a. Abundance of HNA and LNA bacteria co-varied positively in all regions, although HNA bacteria were more responsive to high phytoplankton biomass in shelf waters than in slope and basin waters. Since LNA cell abundance in general showed responses similar to those of HNA cell abundance to changes in phytoplankton biomass, our data do not support the hypothesis that HNA cells are the sole active component of marine bacterioplankton.     

Dissolved and particulate organic carbon and nitrogen in the Northwestern Mediterranean

The distribution of dissolved organic carbon (DOC) and nitrogen (DON) and particulate organic carbon (POC) and nitrogen (PON) was studied on a transect perpendicular to the Catalan coast in the NW Mediterranean in June 1995. The transect covered a hydrographically diverse zone, including coastal waters and two frontal structures (the Catalan and the Balear fronts). The cruise was conducted during the stratified period, characterized by inorganic nutrient depletion in the photic zone and a well established deep chlorophyll a maximum. DOC concentrations were measured using a high-temperature catalytic oxidation method, and DON was determined directly, with an update of the Kjeldahl method, after removal of inorganic nitrogen.The ranges of DOC and DON concentrations were 44–95 μM-C and 2.8–6.2 μM-N. The particulate organic matter ranged between 0.9 and 14.9 μM-C and from 0.1 to 1.7 μM-N. The DOC : DON molar ratio averaged 15.5±0.4, and the mean POC : PON ratio was 8.6±0.6. The distribution of dissolved organic matter (DOM) was inverse to that of the salinity. The highest concentrations of DOM were found in coastal waters and in the stations affected by the Catalan front, located at the continental shelf break.It was estimated that recalcitrant DOM constituted 67% of the DOM pool in the upper 50 m. The data suggest that accumulation of DOC due to the decoupling of production and consumption may occur in the NW Mediterranean during stratification and that the organic matter exported from the photic layer is dominated by C-rich material.     

Zooplankton associated with the oxygen minimum zone system in the northern upwelling region of Chile during March 2000

Zooplankton in the coastal upwelling region off northern Chile may play a significant biogeochemical role by promoting carbon flux into the subsurface OMZ (oxygen minimum zone). This work identifies the dominant zooplankton species inhabiting the area influenced by the OMZ in March 2000 off Iquique (20°S, northern Chile). Abundance and vertical distribution studies revealed 17 copepod and 9 euphausiid species distributed between the surface and 600 m at four stations sampled both by day and by night. Some abundant species remained in the well-oxygenated upper layer (30 m), with no evidence of diel vertical migration, apparently restricted by a shallow (40–60 m) oxycline. Other species, however, were found closely associated with the OMZ. The large-sized copepod Eucalanus inermis was found below the oxycline and performed diel vertical migrations into the OMZ, whereas the very abundant Euphausia mucronata performed extensive diel vertical migrations between the surface waters and the core of the OMZ (200 m), even crossing it. A complete assessment of copepods and euphausiids revealed that the whole sampled water column (0–600 m) is occupied by distinct species having well-defined habitats, some of them within the OMZ. Ontogenetic migrations were evident in Eucalanidae and E. mucronata. Estimates of species biomass showed a substantial (>75% of total zooplankton biomass) daily exchange of C between the photic layer and the OMZ. Both E. inermis and E. mucronata can actively exchange about 37.8 g C m⁻² d⁻¹ between the upper well-oxygenated (0–60 m) layer and the deeper (60–600 m) OMZ layer. This migrant biomass may contribute about 7.2 g C m⁻² d⁻¹ to the OMZ system through respiration, mortality, and production of fecal pellets within the OMZ. This movement of zooplankton in and out of the OMZ, mainly as a result of the migratory behavior of E. mucronata, suggests a very efficient mechanism for introducing large amounts of freshly produced carbon into the OMZ system and should, therefore, be considered when establishing C budgets for coastal upwelling systems.     

Modelling phytoplankton deposition to Chesapeake Bay sediments during winter–spring: interannual variability in relation to river flow

The often-rapid deposition of phytoplankton to sediments at the end of the spring phytoplankton bloom is an important component of benthic–pelagic coupling in temperate and high latitude estuaries and other aquatic systems. However, quantifying the flux is difficult, particularly in spatially heterogeneous environments. Surficial sediment chlorophyll-a, which can be measured quickly at many locations, has been used effectively by previous studies as an indicator of phytoplankton deposition to estuarine sediments. In this study, surficial sediment chlorophyll-a was quantified in late spring at 20–50 locations throughout Chesapeake Bay for 8 years (1993–2000). A model was developed to estimate chlorophyll-a deposition to sediments using these measurements, while accounting for chlorophyll-a degradation during the time between deposition and sampling. Carbon flux was derived from these estimates via C:chl-a = 75.Bay-wide, the accumulation of chlorophyll-a on sediments by late spring averaged 171 mg m⁻², from which the chlorophyll-a and carbon sinking fluxes, respectively, were estimated to be 353 mg m⁻² and 26.5 gC m⁻². These deposition estimates were ∼50% of estimates based on a sediment trap study in the mid-Bay. During 1993–2000, the highest average chlorophyll-a flux was in the mid-Bay (248 mg m⁻²), while the lowest was in the lower Bay (191 mg m⁻²). Winter–spring average river flow was positively correlated with phytoplankton biomass in the lower Bay water column, while phytoplankton biomass in that same region of the Bay was correlated with increased chlorophyll-a deposition to sediments. Responses in other regions of the Bay were less clear and suggested that the concept that nutrient enrichment in high flow years leads to greater phytoplankton deposition to sediments may be an oversimplification. A comparison of the carbon flux associated with the deposition of the spring bloom with annual benthic carbon budgets indicated that the spring bloom did not contribute a disproportionately large fraction of annual carbon inputs to Chesapeake Bay sediments. Regional patterns in chlorophyll-a deposition did not correspond with the strong regional patterns that have been found for plankton net community metabolism during spring.     

Seasonal influences on size-fractionated chlorophyll a concentrations and primary production in the north-west Indian Ocean

Chlorophyll a (chl a) concentrations and primary production by the 0.2–2, 2–18 and >18 μm phytoplankton size-fractions were estimated along a transect in the NW Indian Ocean extending from the coast of Oman to 8°N 68°E during the late SW monsoon and autumn intermonsoonal seasons in 1994. Primary production was estimated using the ¹⁴C technique with either in situ or simulated in situ incubations. During the late monsoon season, maximal chl a and production values were recorded in the coastal upwelling zone with values of 69 mg m^-2 and 3800 mg C m^-2 d^-1, respectively. The maxima, which were distributed patchily in this region, were dominated by the >18 μm size-fraction. Over the remainder of the transect chl a concentrations and production averaged 30 mg m^-2 and 1500 mg C m^-2 d^-1, respectively, with approximately equal contributions by the three size-fractions in the case of chl a at the majority of stations, but in general, with a maximum in production in the 0.2–2 μm fraction. Immediately following cessation of the SW monsoon wind, chl a and production values over the northern part of the transect decreased to values similar to those over the southern part of the transect at the time of the SW monsoon, with the contributions by the three size-fractions being approximately equal. During the following intermonsoonal season, both chl a concentrations and production across the section were dominated by the 0.2–2 μm size-fraction, with average chl a and production values of the order of 20 mg m^-2 and 750 mg C m^-2 d^-1, respectively. Considerable variation in production values, however, was exhibited across the transect. A clearly defined subsurface chl a maximum was only recorded at the southernmost stations of the transect in oligotrophic waters: the feature did not develop universally across the transect during the intermonsoon.     

Cross-slope zonation of erosion and deposition in the Faeroe-Shetland Channel,North Atlantic Ocean

Boundary currents and internal waves determine cross-slope zonation of erosion and deposition in the Faeroe-Shetland Channel. Currents were measured at 8 and 34–50 m above the bottom at three mooring sites (502, 595 and 708 m depth) for 14 days. The structure of the water column was evaluated from CTD sections, and included nepheloid layers and particulate matter concentrations. Indicators for recent deposition in the sediment (organic carbon, phytopigments, ²¹⁰Pb) were measured at eight stations across the slope. Strong near-bottom currents at the upper slope sustain down-slope particle transport in a benthic nepheloid layer, which is eroded under the influence of critically reflecting M₂ internal tidal waves at 350–550 m, where the major pycnocline meets the sloping bottom. Beam attenuation profiles confirmed the presence of intermediate nepheloid layers intruding into the Channel along the major pycnocline, and elevated concentrations of particulate matter and chlorophyll-a were measured at this depth. Near-bottom currents decreased with depth, thus allowing particle deposition down the slope. Inventories of excess ²¹⁰Pb activity in the sediment deeper than 600 m were higher than what was expected on the basis of atmospheric input of ²¹⁰Pb and production in the water column, thus indicating additional lateral inputs. Simple calculations showed that off-slope input of particles from areas shallower than 600 m may be responsible for the enhanced deposition at greater depths.     

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”.     

Spatial and temporal variability of POC in the northeast Subarctic Pacific

Particulate organic carbon (POC) concentrations from 0 to 1000 m were quantified in size-fractionated particulate matter samples obtained by the multiple unit large volume in situ filtration system (MULVFS) in 1996 and 1997 along the 1600 km long “line P” transect from continental slope waters near southern Vancouver Island to Ocean Station PAPA (OSP, 50°N, 145°W). Regression of in situ POC vs. beam attenuation coefficient, c, from a simultaneously deployed 1-m pathlength SeaTech transmissometer gave slope, intercept and r² values of 6.15±0.19×10⁻⁵ m⁻¹ (nmol C l⁻¹)⁻¹, 0.363±0.003 m⁻¹, and 0.951 (n=145), respectively. This result agreed within several percent of calibrations obtained from two 2600-km-long transects of the equatorial Pacific in 1992 (Bishop, 1999). Data from other, more frequently deployed transmissometers were standardized against the 1-m instrument, and the combined optical data set was used to document POC variability at finer spatial and temporal scales than could be sampled directly using either conventional water bottle casts or MULVFS. Published bottle POC vs. c relationships show much more variability and remain problematic. Along the line P transect in the salinity-stratified upper 100 m, POC isolines shoaled from winter to summer in concert with seasonal stratification. At the same time, POC was progressively enriched in subeuphotic zone waters to depths greater than 500 m. Near-surface POC fields sampled in the winter time showed strong temporal POC variability over time scales of days as well as between years. POC concentrations at OSP in February 1996 were higher than those found at any other time of year. Less variability was found along line P in other seasons. In May 1996, kilometer-scale spatial variability of POC at OSP was small; dawn vs. dusk variations of c were used to calculate 0–100 m POC turnover times shorter than 6 d. Calculations also suggest that 25–50% of primary productivity was expressed as dissolved organic carbon at OSP in May 1996.     

Arsenic and antimony species in surface transects and depth profiles across a frontal zone: The Chatham Rise,New Zealand

Measurements of total dissolved arsenic (As(III+V)) and antimony (Sb(III+V) and their simple methylated species are presented for samples collected from three vertical profiles and along three surface transects in the Chatham Rise region, east of New Zealand. As(III+V) concentrations showed a slight increase with depth (16–17 nM at 25 m to 20 nM at 100 m) whereas Sb(III+V) concentrations were conservative with depth (1.02–1.12 nM). Along the three surface water transects, As(III+V) and Sb(III+V) concentrations showed little variation, with average concentrations of 18±2 and 0.99±0.05 nM, respectively. Inorganic arsenic was not correlated with orthophosphate (r²=0.01). Monomethyl- and dimethyl-arsenic (MMAs, DMAs) concentrations (0.04–0.01 and 0.65–0.07 nM, respectively) decreased with depth, suggesting surface water production by biota and degradation at depth. Along the Chatham Rise transect, DMAs concentrations increased on the Rise (0.65 nM maximum) compared to waters north and south of the Rise (∼0.22 nM). Fluctuation in MMAs concentrations were also seen for water samples collected on the Chatham Rise. Monomethyl-, dimethyl- and trimethyl-antimony (MMSb, DMSb, TMSb) species were detected in water samples collected along all the three surface water transects suggesting surface water production by biota. Concentrations of MMSb, DMSb and TMSb in water samples were fairly constant along all the three surface transects (0.06–0.07, 0.015–0.025 and 0.005–0.015 nM, respectively), showing no significant enrichment on the Chatham Rise. These arsenic and antimony results support the current global view that inorganic As and Sb are conservative and the methyl species are of biological origin.     

Nutrient gradients in the western North Atlantic Ocean: Relationship to microbial community structure and comparison to patterns in the Pacific Ocean

Studies of nitrogen and phosphorus dynamics in the oligotrophic surface waters of the western North Atlantic Ocean have been constrained because ambient concentrations are typically at or below the detection limits of standard colorometric methods, except during periods of deep vertical mixing. Here we report the application of high-sensitivity analytical methods—determinations of nitrate plus nitrite (N+N) by chemiluminescence and soluble reactive phosphorus (SRP) by the magnesium induced co-precipitation (MAGIC) protocol—to surface waters along a transect from the Sargasso Sea at 26°N through the Gulf Stream at 37°N, including sampling at the JGOFS Bermuda Atlantic Time-series Study (BATS) station. The results were compared with data from the BATS program, and the HOT station in the Pacific Ocean, permitting cross-ecosystem comparisons. Microbial populations were analyzed along the transect, and an attempt was made to interpret their distributions in the context of the measured nutrient concentrations.Surface concentrations of N+N and SRP during the March 1998 transect separated into 3 distinct regions, with the boundaries corresponding roughly to the locations of the BATS station (∼31°N) and the Gulf Stream (∼37°N). Although N+N and SRP co-varied, the [N+N] : [SRP] molar ratios increased systematically from ∼1 to 10 in the southern segment, remained relatively constant at ∼40–50 between 31°N and 37°N, then decreased again systematically to ratios <10 north of the Gulf Stream. Dissolved organic N (DON) and P (DOP) dominated (⩾90%) the total dissolved N (TDN) and P (TDP) pools except in the northern portion of the transect. The [DON] : [DOP] molar ratios were relatively invariant (∼30–60) across the entire transect.Heterotrophic prokaryotes (operationally defined as “bacteria”), Prochlorococcus, Synechococcus, ultra- and nanophytoplankton, cryptophytes, and coccolithophores were enumerated by flow cytometry. The abundance of bacteria was well correlated with the concentration of SRP, and that of the ultra- and nanophytoplankton was well correlated with the concentration of N+N. The only group whose concentration was correlated with temperature was Prochlorococcus, and its abundance was unrelated to the concentrations of nutrients measured at the surface.We combined our transect results with time-series measurements from the BATS site and data from select depth profiles, and contrasted these North Atlantic data sets with time-series of N and P nutrient measurements from a station in the North Pacific subtropical gyre near Hawaii [Hawaii Ocean Time-series (HOT) site]. Two prominent differences are readily observed from this comparison. The [N+N] : [SRP] molar ratios are much less than 16 : 1 during stratified periods in surface waters at the BATS site, as is the case at the HOT site year round. However, following deep winter mixing, this ratio is much higher than 16 : 1 at BATS. Also, SRP concentrations in the upper 100 m at BATS fall in the range 1–10 nM during stratified periods, which is at least one order of magnitude lower than at the HOT site. That two ecosystems with comparable rates of primary and export production would differ so dramatically in their nutrient dynamics is intriguing, and highlights the need for detailed cross ecosystem comparisons.     

Planktic foraminiferal production stimulated by chlorophyll redistribution and entrainment of nutrients

During September and October 1996 planktic foraminifers and pteropods were sampled from the upper 2500 m of the water column in the BIOTRANS area (47°N, 20°W), eastern North Atlantic, as part of the JGOFS program. Hydrography, chlorophyll fluorescence, and nutrient content were recorded at high spatial and temporal resolution providing detailed information about the transition time between summer and fall. At the beginning of the cruise a shallow pycnocline was present and oligotrophic conditions prevailed. Over the course of the cruise, the mixed layer depth increased and surface water temperature decreased by 1.5°C. Both chlorophyll-a dispersed in the upper 50 m by vertical mixing and chlorophyll-a concentrations at the sea surface increased. The nitracline shoaled and nutrient enriched waters were entrained into the mixed layer. Planktic foraminifers and pteropods closely reflected the changes in the hydrography by increased growth rates and changes in species composition. Three main groups of planktic foraminiferal species were recognized: (1) a temperate and low-productivity group dominated by Neogloboquadrina incompta characterized the shallow mixed layer depths. (2) A temperate and high-productivity group dominated by Globigerina bulloides characterized the period with wind-induced dispersal of chlorophyll-a and entrainment of nutrient-enriched waters. (3) A warm water group containing Globigerinoides sacculifer, Orbulina universa, Globigerinoides ruber (white), and Globigerinella siphonifera was most common during the first days of sampling. Synchronous with the hydrographic change from summer to fall, planktic foraminiferal and pteropod growth was stimulated by redistribution of chlorophyll-a and entrainment of nutrient-enriched waters into the mixed layer. In addition, the seasonal change in the eastern North Atlantic resulted in a transition of the epipelagic faunal composition and an increased calcareous particle flux, which could be used to trace seasonality in fossil assemblages and allow for better paleoceanographic interpretation of the boreal Atlantic.     

Variability of chlorophyll and primary production in the Eastern North Atlantic Subtropical Gyre: potential factors affecting phytoplankton activity

Size-fractionated chlorophyll-a and carbon incorporation rates were determined on a series of 13 cruises carried out from 1992 to 2001with the aim of investigating the patterns and causes of variability in phytoplankton chlorophyll and production in the Eastern North Atlantic Subtropical Gyral Province (NASE). Averaged (±SE) integrated chlorophyll-a concentration and primary production rate were 17±1 mg m⁻² and 253±22 mg C m⁻² d⁻¹. Small-sized cells (<2 μm) formed the bulk of phytoplankton biomass (71%) and accounted for 54% of total primary production. A clear latitudinal gradient in these variables was not detected. By contrast, large seasonal variability was detected in terms of primary production, although integrated phytoplankton biomass, as estimated from chlorophyll-a concentration, remained rather constant and did not display significant changes with time. Variability in primary production (PP) was related mainly to variability in surface temperature and surface chlorophyll-a concentration. The control exerted by surface temperature was related to nutrient availability. By contrary, euphotic-zone depth, depth of maximum concentration of chlorophyll-a and integrated chlorophyll-a did not contribute significantly to the high variability in primary production observed in this oligotrophic region.     

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.     

Distribution and diversity of open-ocean,near-bottom macroplankton in the western Mediterranean: Analysis at different spatio-temporal scales

We analyzed the distribution, diversity, and composition of western Mediterranean macroplankton (excluding gelatinous taxa) in the water column over depths of ca. 550–850 m, with special attention to near-bottom (0–1.5 and ca. 5–77 m above the bottom, mab) levels, and including data from three areas (off the coasts of Catalonia, and to the NW, and SE of Mallorca, Balearic Islands) in the period 1991–2008. Spatio-temporal changes in macroplankton abundance were evaluated as follows: (i) by seasonal sampling in 2007 off the Catalonian coast, (ii) by comparing Catalonian and Balearic Island slopes, and (iii) by comparing a fixed station on the Catalonian slope (at 550–800 m depth) at decadal (1991/1992–2007/2008) time scales. Diversity (in terms of species richness, S) was greater (i) at ca. 5–77 mab than at 0–1.5 mab, (ii) over the insular slopes of the Balearic Island (around Mallorca) than over the mainland Catalonian slopes, and (iii) in the period 1991/1992 than in 2007, likely related to higher values of the North Atlantic Oscillation (NAO) index in 1991/1992. In most analyses species composition was strongly influenced by the degree of stratification and homogenization of the water column in summer–autumn and winter–spring respectively and by location (longitude). Changes consisted mainly of higher density of macroplankton (e.g. abundance of the dominant euphausiids Nematoscelis megalops, Meganyctiphanes norvegica and Euphausia krohni and of the fish Cyclothone braueri) between June and October, parallel to an increase in the T and S close to the bottom. This coincided with changes in the flow of Levantine intermediate water (LIW) in the area. Aggregation of adult forms of the dominant species close to the bottom in summer–autumn could be favored because summer is the period of highest density of food – copepods, mainly Calanus helgolandicus – near the bottom off the Catalan slope. The formation of a thermocline in the water column and the reinforcement of the permanent thermohaline front at the shelf-slope break during summer at ca. 400 m in the Balearic Basin may also enhance this tendency toward greater aggregation of deep macroplankton under stratified water column conditions.     

Observations on depth distribution,diversity and abundance of pelagic nemerteans from the Pacific Ocean off California and Hawaii

Meso- and bathypelagic nemerteans were collected during five cruises west of Pt. Conception, California, and one west of Oahu, Hawaii, from depths of 0–3800 m, using a 10 m² Tucker Trawl modified with a thermally protected closing cod end, as well as MOCNESS-10 and MOCNESS-1D trawls. In this study, about 53 putative species were collected off California and about 23 off Hawaii, with little taxonomic overlap between the two groups, as compared to 16 and 0 species previously known from these regions. Numbers of nemertean individuals and species were greatest between 625 and 1750 m off California, and 1300–2500 m off Hawaii. Abundance of nemerteans off California was about 10 times that off Hawaii. The Shannon– Wiener diversity index (H′) was higher for Hawaii than for California due to numerical dominance by two species, Phallonemertes cf murrayi and Nectonemertes cf mirabilis, off California and a more even species distribution off Hawaii. Neither of the two dominant California species showed evidence of nocturnal vertical migration to shallow water. This is the first report of a Phallonemertes species from the Pacific and the first report of pelagic nemerteans from waters off Hawaii.     

Organic complexation of iron in the Southern Ocean

The chemical speciation of iron was determined in the Southern Ocean along a transect from 48 to 70°S at 20°E. Dissolved iron concentrations were low at 0.1–0.6 nM, with average concentrations of 0.25±0.13 nM. Organic iron complexing ligands were found to occur in excess of the dissolved iron concentration at 0.72±0.23 nM (equivalent to an excess of 0.5 nM), with a complex stability of log K_FeL′=22.1±0.5 (on the basis of Fe³⁺ and L′). Ligand concentrations were higher in the upper water column (top 200 m) suggesting in situ production by microorganisms, and less at the surface consistent with photochemical breakdown. Our data are consistent with the presence of stable organic iron-complexing ligands in deep global ocean waters at a background level of ∼0.7 nM. It has been suggested that this might help stabilise iron at levels of ∼0.7 nM in deep ocean waters. However, much lower iron concentrations in the waters of the Southern Ocean suggest that these ligands do not prevent the removal of iron (by scavenging or biological uptake) to well below the concentration of these ligands. Scavenging reactions are probably inhibited by such ligand competition, so it is likely that biological uptake is the chief cause for the further removal of iron to these low levels in waters that suffer from very low iron inputs.     

Behaviors of dissolved and particulate Co,Ni, Cu,Zn, Cd and Pb during a mesoscale Fe-enrichment experiment (SEEDS II) in the western North Pacific

During mesoscale Fe enrichment (SEEDS II) in the western North Pacific ocean, we investigated dissolved and particulate Co, Ni, Cu, Zn, Cd and Pb in seawater from both field observation and shipboard bottle incubation of a natural phytoplankton assemblage with Fe addition. Before the Fe enrichment, strong correlations between dissolved trace metals (Ni, Zn and Cd) and PO₄³⁻, and between particulate trace metals (Ni, Zn and Cd) and chlorophyll-a were obtained, suggesting that biogeochemical cycles mainly control the distributions of Ni, Zn and Cd in the study area. Average concentrations of dissolved Co, Ni, Cu, Zn, Cd and Pb in the surface mixed layer (0–20 m) were 70 pM, 4.9, 2.1, 1.6, 0.48 nM and 52 pM, respectively, and those for the particulate species were 1.7 pM, 0.052, 0.094, 0.46, 0.037 nM and 5.2 pM, respectively. After Fe enrichment, chlorophyll-a increased 3 fold (up to 3 μg L⁻¹) during developing phases of the bloom (<12 days). Mesozooplankton biomass also increased. Particulate Co, Ni, Cu and Cd inside the patch hinted at an increase in the concentrations, but there were no analytically significant differences between concentrations inside and outside the patch. The bottle incubation with Fe addition (1 nM) showed an increase in chlorophyll-a (8.9 μg L⁻¹) and raised the particulate fraction up to 3–45% for all the metals, accompanying changes in Si/P, Zn/P and Cd/P. These results suggest that Fe addition lead to changes in biogeochemical cycling of trace metals. The comparison between the mesoscale Fe enrichment and the bottle incubation experiment suggests that although Fe was a limiting factor for the growth of phytoplankton, the enhanced biomass of mesozooplankton also limited the growth of phytoplankton and the transformation of trace metal speciation during the mesoscale Fe enrichment. Sediment trap data and the elemental ratios taken up by phytoplankton suggest that export loss was another reason that no detectable change in the concentrations of particulate trace metals was observed during the mesoscale Fe enrichment.     

High short-term variability of CO2 fluxes during an upwelling event off the Chilean coast at 30°S

pH and alkalinity measurements from a coastal upwelling area located near 30°S (Coquimbo, Chile), are used to describe the short-term variations of CO₂ air–sea exchanges over a period of one week in summer 1996. A 180 km ocean–coastal transect, together with two almost-synoptic grid surveys off Coquimbo covering approximate 2500 km² each, showed that during and immediately after a 4 day long southwesterly wind event (24–28 January) a large area of cold surface water (≈14°C), highly supersaturated in CO₂ (fCO₂ up to 900 μatm), was located near the coast. Three days after the end of the event, the second grid survey showed that in most of the study area the surface temperature and pH had increased significantly (by 1–3°C and 0.05–0.2, respectively), and that the surface water was no longer supersaturated in CO₂. The CO₂-supersaturated water observed in the first grid survey was identified as upwelled subsurface equatorial water, a water mass with its core at about 200 m depth: the depth from which the water upwells is a major determinant of the surface water fCO₂. Integrated C fluxes within a 20 km wide coastal strip (1900 km²) indicate a strong outgassing of CO₂ from the ocean under upwelling conditions (Grid 1; 121 t C day^-1), while the net C exchange was directed to the ocean during the relaxation period (Grid 2; 19 t C day^-1). Estimates of CO₂ fluxes in upwelling areas based on surface water fCO₂ measurements must therefore take into account these short-term variations: reliance on longer-term averages and interpolation will lead to erroneous results.     

Iron fertilization of Trichodesmium off the west coast of Barbados: A one-dimensional numerical model

A one-dimensional quasi-Eulerian model, describing the biological and chemical interactions of autotrophic and heterotrophic plankton populations within the upper 100 m of the water column, was used to explain the relative impact of iron fertilization from Saharan dust on phosphorus cycling by the cyanobacterium, Trichodesmium spp. We examined the Caribbean surface waters off the west coast of Barbados using seasonal cases of dust delivery (summer peak) in relation to periods of elevated phosphorus stocks found in episodic pulses of Amazon River water advected past Barbados by the Guiana Current. The additional iron and phosphorus supplied by fluvial sources, available after biological depletion of nitrogen, could alleviate the growth limitation of Trichodesmium far downstream of the river plumes. The model results were compared with measurements made of Trichodesmium stocks 8 km off the west coast of Barbados. Modeled concentrations within the low-salinity Amazon plumes reached ∼7.4 mg chl m⁻², as seen at Barbados. In the absence of the low-salinity signal, the Trichodesmium biomass shifted to phosphorus limitation with little response to iron fertilization. This indicates that Trichodesmium stocks may indeed be mainly a function of phosphorus availability within the upper water column. Therefore, the supply of phosphorus from subtropical/tropical rivers, such as the Amazon and Orinoco, limits the potential cyanophyte response, despite adequate atmospheric iron delivery. This alters our view not only of the western North Atlantic nitrogen budget, but potentially several regions downstream of low N:P river systems.