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

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

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.     

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.     

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”.     

Wave climate variability in the North-East Atlantic Ocean over the last six decades

Ocean surface gravity waves play a major role in many engineering and environmental problems, both in the open ocean and in coastal zones. Therefore, it is essential to improve our knowledge on spatial and temporal variability of wave climate. This study aims at investigating this variability in the North-East Atlantic Ocean (25°W–0°W and 30°N–60° N), using a 57-year hindcast (1953–2009) obtained with a spectral wave model forced with reanalysis wind fields. The hindcast analysis reveals firstly strong seasonal fluctuations of wave climate, with winters characterized by large and long-period waves of mean direction spreading from south-west to north-west, and summers characterized by smaller and shorter-period waves originating from norther directions. From northern (55°N) to southern (35°N) latitudes, the significant wave height (Hs) decreases by roughly 40%, the mean wave direction (Mwd) rotates clockwise by about 25% while the peak period (Tp) only grows by 5%. These three parameters also exhibit a strong inter-annual variability, particularly when winter-means (from 1st of December to 1st of April) are considered. Linear trend analysis over the studied period shows spatially variable long-term trends, with a significant increase of Hs (up to 0.02 m yr^?1) and a counterclockwise shift of Mwd (up to ?0.1° yr^?1) at northern latitude, contrasting with a fairly constant trend for Hs and a clockwise shift of Mwd (up to +0.15° yr^?1) at southern latitudes. Long-term trends of Tp are less significant, with still a slight increase in the north-eastern part of the study area (up to +0.01 s yr^?1). Eventually, a comparison between the inter-annual variability of the winter-means of the three selected wave parameters and the North Atlantic Oscillation (NAO) reveals: (1) a strong positive correlation between Hs and the NAO index at northern latitudes (correlation coefficient up to R = 0.91) and a significant negative correlation at southern latitudes (up to R = ?0.6); (2) no significant correlation for Mwd north of 40°N and a clear positive correlation southward of 40°N (up to R = 0.8) and (3) a northward increasing correlation for Tp (up to R = 0.8). Long-term trends for Hs, Mwd and Tp are finally explained by a significant increase in the NAO index over the studied period.     

Vertical distributions of dissolved organic carbon and nitrogen in the Southern Ocean

Detailed vertical profiles of dissolved organic carbon (DOC), nitrogen (DON) and DOC/DON ratios in the Australian sector of the Southern Ocean (56°–65°S around 140°W transect) were obtained by a high-temperature catalytic oxidation method with a modified Shimadzu TOC-5000 unit. The simultaneous analyses of DOC and total dissolved nitrogen (TDN) gave high precision results (±0.7 μMC and ±0.3 μMN). In surface layers, DOC concentrations were lower (45–55 μM) than those generally obtained from other oceanic environments (60–90 μM). The surface concentrations of DON varied from 4 to 9 μM, but within the range generally reported for other ocean regions. Surface excesses of DOC and DON were calculated against nearly constant subsurface concentrations. A consistent contribution of low C/N ratio (2.7–5.0) was found in the mid-surface layer (30–75 m), suggesting more extensive degradation of carbon-enriched materials and/or enhanced supply of nitrogen-enriched ones.     

Atmospheric iron deposition and sea-surface dissolved iron concentrations in the eastern Atlantic Ocean

Atmospheric iron and underway sea-surface dissolved (<0.2 μm) iron (DFe) concentrations were investigated along a north–south transect in the eastern Atlantic Ocean (27°N/16°W–19°S/5°E). Fe concentrations in aerosols and dry deposition fluxes of soluble Fe were at least two orders of magnitude higher in the Saharan dust plume than at the equator or at the extreme south of the transect. A weaker source of atmospheric Fe was also observed in the South Atlantic, possibly originating in southern Africa via the north-easterly outflow of the Angolan plume. Estimations of total atmospheric deposition fluxes (dry plus wet) of soluble Fe suggested that wet deposition dominated in the intertropical convergence zone, due to the very high amount of precipitation and to the fact that a substantial part of Fe was delivered in dissolved form. On the other hand, dry deposition dominated in the other regions of the transect (73–97%), where rainfall rates were much lower. Underway sea-surface DFe concentrations ranged 0.02–1.1 nM. Such low values (0.02 nM) are reported for the first time in the Atlantic Ocean and may be (co)-limiting for primary production. A significant correlation (Spearman's rho=0.862, p<0.01) was observed between mean DFe concentrations and total atmospheric deposition fluxes, confirming the importance of atmospheric deposition on the iron cycle in the Atlantic. Residence time of DFe in the surface waters relative to atmospheric inputs were estimated in the northern part of our study area (17±8 to 28±16 d). These values confirmed the rapid removal of Fe from the surface waters, possibly by colloidal aggregation.     

Selective depressions of surface silicic acid within cyclonic mesoscale eddies in the oligotrophic western North Pacific

To reveal spatial dynamics of silicic acid [Si(OH)₄] in the poorly sampled oligotrophic western North Pacific, we investigated the surface distribution of Si(OH)₄ and associated biogeochemical parameters by using an underway survey system with a highly sensitive nutrient analyzer along the 138°E transect (between 30 and 34°N) and the 155°E transect (between 10 and 35°N) during the summers of 2007 and 2008. Surface Si(OH)₄ concentrations ranged from the detection limit (11 nmol L⁻¹) to 2462 nmol L⁻¹. High Si(OH)₄ concentrations (>1000 nmol L⁻¹) and dynamic fluctuations were generally observed north of 23°N, while consistently stable low concentrations of 415–751 nmol L⁻¹ were observed south of 23°N. Surface nitrate+nitrite (N+N) and phosphate (PO₄³⁻) were typically depleted to <20 nmol L⁻¹, except for PO₄³⁻ in the area south of 16°N. The majority of the study area was characterized by high-Si(OH)₄ and low-N+N and PO₄³⁻. However, submesoscale/mesoscale depressions of Si(OH)₄ were locally observed in the cyclonic eddy fields north of 23°N. Among a total of six Si(OH)₄ depressions within the eddies, a complete Si(OH)₄ depletion (<11 nmol L⁻¹) was observed on the cyclonic side near the Kuroshio axis (33.1°N, 138°E). This depletion was closely coupled with a diatom bloom, suggesting that Si(OH)₄ was exhausted by diatoms. All of the Si(OH)₄ depressions were selective and not accompanied by local depressions of N+N and PO₄³⁻. This unique phenomenon might be driven by biogeochemical processes such as selective Si export (Si pump), anomalous Si uptake associated with diatom physiology, and/or Si uptake supported by N₂ fixation.     

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

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

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

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

Distributions of particulate and dissolved organic and inorganic phosphorus in North Pacific surface waters

Several operationally defined fractions of phosphorus (P) were measured along a surface water transect in the North Pacific. The P content in all fractions was found to increase northward from the edge of the subtropical to the subarctic region. Particulate organic P (POP) concentrations increased from 9 to 110 nmol L^− 1, whereas the particulate inorganic P (PIP) concentrations increased from 1 to 13 nmol L^− 1. A significant correlation between POP, PIP and chlorophyll a suggested that these P pools are associated directly or indirectly with phytoplankton cells. PIP comprised 10–20% of the total particulate P pool across the transect, indicating it is an important component of the marine P cycle in this region. Dissolved non-reactive P (assumed to consist predominantly of non-reactive organic P compounds, thus referred to as DOP) concentration increased from 0.10 to 0.22 μmol L^− 1, whereas soluble reactive P (SRP) concentration increased from 0.01 to 1.42 μmol L^− 1 along the transect. The proportion of DOP and SRP varied widely, with a large proportion of DOP in areas with low total dissolved P concentrations in lower latitudes and a large proportion of SRP in areas with high total dissolved P concentrations in higher latitudes. High demand for DOP in the lower latitudinal region would diminish the concentration of this pool relative to higher latitudinal regions where SRP is more abundant and would be preferentially utilized. The availability of SRP could have a significant impact on the concentration and probably on the composition of DOP. We show that P fractionation provides an important insight for discussing the marine P cycle.     

Direct observation of subtropical mode water circulation in the western North Atlantic Ocean

Eighteen Degree Water (EDW) is the dominant subtropical mode water of the North Atlantic subtropical gyre and is hypothesized as an interannual reservoir of anomalous heat, nutrients and CO₂. Although isolated beneath the stratified upper-ocean at the end of each winter, EDW may re-emerge in subsequent years to influence mixed layer properties and consequently air–sea interaction and primary productivity. Here we report on recent quasi-Lagrangian measurements of EDW circulation and stratification in the western subtropical gyre using an array of acoustically-tracked, isotherm-following, bobbing profiling floats programmed to track and intensively sample the vertically homogenized EDW layer and directly measure velocity on the 18.5 °C isothermal surface.The majority of the CLIVAR Mode Water Dynamics Experiment (CLIMODE) bobbers drifted within the subtropical gyre for 2.5–3.5 years, many exhibiting complex looping patterns indicative of an energetic eddy field. Bobber-derived Lagrangian integral time and length scales (3 days, 68 km) associated with motion on 18.5 °C were consistent with previous measurements in the Gulf Stream extension region and fall between previous estimates at the ocean surface and thermocline depth. Several bobbers provided evidence of long-lived submesoscale coherent vortices associated with substantial EDW thickness. While the relative importance of such vortices remains to be determined, our observations indicate that these features can have a profound effect on EDW distribution. EDW thickness (defined using a vertical temperature gradient criterion) exhibits seasonal changes in opposition to a layer bounded by the 17 °C and 19 °C isotherms. In particular, EDW thickness is generally greatest in winter (as a result of buoyancy-forced convection), while the 17°–19 °C layer is thickest in summer consistent with seasonal Ekman pumping. Contrary to previous hypotheses, the bobber data suggest that a substantial fraction of subducted EDW is isolated from the atmosphere for periods of less than 24 months. Seasonal-to-biennial re-emergence (principally within the recirculation region south of the Gulf Stream) appears to be a common scenario which should be considered when assessing the climatic and biogeochemical consequences of EDW.     

Year-long float trajectories in the Labrador Sea Water of the eastern North Atlantic Ocean

Year-long Lagrangian trajectories within the Labrador Sea Water of the eastern North Atlantic Ocean are analysed for basic flow statistics. Root-mean-square velocities at 1750 m depth are about 2 cm/s, except within the North Atlantic Current, where they are twice as large. These values are consistent with previous Eulerian measurements and extend those results to a much larger domain of the eastern basin. Mean flow estimates in boxes large enough to contain about 1 float-year of data indicate that Labrador Sea Water, having crossed the Mid- Atlantic Ridge (not resolved) near 50–55^°N, presumably with the North Atlantic Current, partially recirculates to the north in the subpolar gyre, as well as entering the subtropical gyre and continuing south and west. The circulation of this water mass, as defined by the 1 yr average velocities, is stronger than traditional models of deep circulation would suggest, with an interior flow of roughly 1 cm/s. Mean speeds up to 3 cm/s were observed, with the highest values near the Azores Plateau. North of 45°N–55°N, mean eastward speeds closer to 0.2 cm/s were observed. Wind-generated barotropic fluctuations may be responsible for some part of the transport at this depth.     

The seasonal succession of zooplankton in the Southern Ocean south of Australia,part I: The seasonal ice zone

Between November 2001 and March 2002 an Australian/Japanese collaborative study completed six passes of a transect line in the Seasonal-Ice Zone (south of 62°S) along 140°E. Zooplankton samples were collected with a NORPAC net on 22–28 November, and a Continuous Plankton Recorder on 10–15 January, 11–12 February, 19–22 February, 25–26 February, and 10–11 March. Zooplankton densities were lowest on 22–28 November (ave=61 individuals (ind) m⁻³), when almost the entire transect was covered by sea ice. By 10–15 January sea surface temperature had increased by ∼2 °C across the transect line, and the study area was ice-free. Total zooplankton abundance had increased to maximum levels for the season (ave=1301 ind m⁻³; max=1979 ind m⁻³), dominated by a “Peak Community” comprising Oithona similis, Ctenocalanus citer, Clausocalanus laticeps, foraminiferans, Limacina spp., appendicularians, Rhincalanus gigas and large calanoid copepodites (C1–3). Total densities declined on each subsequent transect, returning to an average of 169 ind m⁻³ on 10–11 March. The seasonal density decline was due to the decline in densities of “Peak Community” taxa, but coincided with the rise of Euphausia superba larvae into the surface waters, increased densities of Salpa thompsoni, and an increased contribution of C4 to adult stages to the populations of Calanoides acutus, Calanus propinquus and Calanus simillimus. The seasonal community succession appeared to be influenced by the low sea ice extent and southward projection of the ACC in this region. The relatively warm ACC waters, together with low krill biomass, favoured high densities of small grazers during the January/February bloom conditions. The persistence of relatively warm surface waters in March and the seasonal decrease in chlorophyll a biomass provided favorable conditions for salps, which were able to penetrate south of the Southern Boundary.     

Seasonality of particle-associated trace element fluxes in the deep northeast Atlantic Ocean

In the framework of the German contribution to the Joint Global Ocean Flux Study (JGOFS), deep-water fluxes of particle-associated trace elements were measured in the northeast Atlantic Ocean. The sinking particles were collected almost continuously from 1992 to 1996 at three time-series stations, L1 (33°N/22°W), L2 (47°N/20°W), and L3 (54°N/21°W), using sediment traps. The focus of the present study is the temporal variability of the particle-associated elemental fluxes of Al, Ca, Cd, Co, Cu, Fe, Mn, Ni, P, Pb, Ti, V, and Zn at a depth of 2000 m.A clear seasonality of the fluxes that persisted for several years was documented for the southernmost station (L1) at stable oligotrophic conditions in the area of the North Atlantic Subtropical Gyre East (NASTE). At L2 and L3, an episodic nature of the elemental fluxes was determined. Mesoscale eddies are known to frequently cause temporal and spatial variability in the flux of biogenic components in that area. These events modified the simple seasonal pattern controlled by the annual cycle at L2, in the North Atlantic Drift Region (NADR), and at L3, which was influenced by the Atlantic Arctic province (ARCT). All stations were characterized by an additional episodic lithogenic atmospheric supply reaching the deep sea.The integrated annual fluxes during the multi-year study revealed similar flux magnitudes for lithogenic elements (Al, Co, Fe, Ti, and V) at L2 and L3 and roughly twofold fluxes at L1. Biogenic elements (Cd, P, and Zn) showed the opposite trend, i.e., two to fourfold higher values at L2 and L3 than at L1. For Mn, Ni, and Cu, the spatial differences were smaller, perhaps because of the intermediate behavior, between lithogenic and biogenic, of these elements. Similarly, among the three study sites, there were no noticeable differences in the total annual flux of Pb.The respective lithogenic fractions of the deep-sea fluxes of Cd, Co, Cu, Mn, Ni, V, and Zn were subtracted based on the amount of Al, with the average composition of the continental crust as reference. This procedure allowed estimation of the labile trace element fraction (TE_exc) of the particles, i.e., TE taken up or scavenged during particle production and sedimentation. The ratios of TE_exc/P clearly demonstrated an enrichment of TE over labile P from biogenic surface material to the deep sea for Zn (factor 4–6), Mn (12–27), Ni (3–5), and Cu (9–25); an intermediate status for Co (0.5–2.2); and depletion for Cd vs. P (0.2–0.4). Surprisingly, the recycling behavior of excess Co was found to be similar to that of P. Hence, Co_exc behaved like a biogenic element; this is in contrast to total Co, which is dominated by the refractory lithogenic fraction.Moreover, it is argued that these excess elemental fluxes caused a loss of the dissolved elements in upper waters, since their transport reaches the deep-sea waters at 2000 m, a depth far below of deep-winter mixing and upwelling. The annual amount of excess TE exported from surface waters was estimated to be 1.3×10⁹ mol Zn y^?1, 4.4×10⁹ mol Mn y^?1, 4.9×10⁸ mol Ni y^?1, 2.2×10⁷ mol Cd y^?1, 7.4×10⁸ mol Cu y^?1, and 2.7×10⁷ mol Co y^?1 for the whole North Atlantic Ocean. Important primary sources that could replenish these losses are the aeolian and fluvial supply processes.     

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

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

Oxygen dynamics in the North Atlantic subtropical gyre

Dissolved oxygen (DO) in the ocean is a tracer for most ocean biogeochemical processes including net community production and remineralization of organic matter which in turn constrains the biological carbon pump. Knowledge of oxygen dynamics in the North Atlantic Ocean is mainly derived from observations at the Bermuda Atlantic Time-series Study (BATS) site located in the western subtropical gyre which may skew our view of the biogeochemistry of the subtropical North Atlantic. This study presents and compares a 15 yr record of DO observations from ESTOC (European Station for Time-Series in the Ocean, Canary Islands) in the eastern subtropical North Atlantic with the 20 yr record at BATS. Our estimate for net community production of oxygen was 2.3±0.4 mol O₂ m⁻² yr⁻¹ and of oxygen consumption was −2.3±0.5 mol O₂ m⁻² yr⁻¹ at ESTOC, and 4 mol O₂ m⁻² yr⁻¹ and −4.4±1 mol m⁻² yr⁻¹ at BATS, respectively. These values were determined by analyzing the time-series using the Discrete Wavelet Transform (DWT) method. These flux values agree with similar estimates from in-situ observational studies but are higher than those from modeling studies. The difference in net oxygen production rates supports previous observations of a lower carbon export in the eastern compared to the western subtropical Atlantic. The inter-annual analysis showed clear annual cycles at BATS whereas longer cycles of nearly 4 years were apparent at ESTOC. The DWT analysis showed trends in DO anomalies dominated by long-term perturbations at a basin scale for the consumption zones at both sites, whereas yearly cycles dominated the production zone at BATS. The long-term perturbations found are likely associated with ventilation of the main thermocline, affecting the consumption and production zones at ESTOC.     

Microzooplankton composition,biomass and grazing rates along the WOCE SR3 line between Tasmania and Antarctica

Microzooplankton species composition and grazing rates on phytoplankton were investigated along a transect between ∼46 and 67°S, and between 140 and 145°E. Experiments were conducted in summer between November 2nd and December 14th in 2001. The structure of the microbial food web changed considerably along the transect and was associated with marked differences in the physical and chemical environment encountered in the different water masses and frontal regions. On average microzooplankton grazing experiments indicated that 91%, 102%, and 157%, (see results) of the phytoplankton production would be grazed in the <200, <20 and <2 μm size fractions, respectively, indicating microzooplankton grazing was potentially constraining phytoplankton populations (<200 μm) along most of the transect. Small ciliates in general and especially oligotrich species declined in importance from the relatively warm, Southern Subtropical Front waters (6.8 μg C/L) to the colder waters of the southern branch of the Polar Front (S-PF), (∼0.5 μg C/L) before increasing again near the Antarctic landmass. Large changes in microzooplankton dominance were observed, with heterotrophic nanoflagellates (HNF), ciliates and larger dinoflagellates having significant biomass in different water masses. HNF were the dominant grazers when chlorophyll a was low in areas such as the Inter-Polar Frontal Zone (IPFZ), while in areas of elevated biomass such as the S-PF and Southern Antarctic Circumpolar Current (SACC), a mix of copepod nauplii and large heterotrophic and mixotrophic dinoflagellates tended to dominate the grazing community. In the S-PF and SACC water masses the tight coupling observed between the microzooplankton grazers and phytoplankton populations over most of the rest of the transect was relaxed. In these regions grazing was low on the >20 μm size fraction of chlorophyll a, which dominated the biomass, while smaller diatoms and nanoplankton in the <20 μm size fraction were still heavily grazed. The lack of grazing pressure on large phytoplankton contributes to this region's potential to export carbon with larger cells known to have higher sinking rates.     

The swarm dynamics of northern krill (Meganyctiphanes norvegica) and pteropods (Cavolinia inflexa) during vertical migration in the Ligurian Sea observed by an acoustic Doppler current profiler

A ship-mounted 153 kHz narrow-band ADCP and 1 m² MOCNESS were deployed between 16 and 24 Sept. 1997 in the Ligurian central zone (∼43°20′N 7°48′E). Results from both instruments showed that the zooplankton community performed vertical migrations that conformed to the classical pattern of ascent at dusk (∼18:30 h) and descent at dawn (∼06:30 h). Depth-discrete net samples between 0 and 500 m showed that the community was dominated by two species, the euphausiid Meganyctiphanes norvegica (Northern krill) and the pteropod Cavolinia inflexa, which migrated in separate discrete bands that were detectable by the ADCP. Information from the ADCP was used to estimate vertical migration speed in two ways: (i) from the trajectory of the back-scattering bands over time and (ii) from the Doppler-shift vertical velocity measured within depth zones at the corresponding time and depth of these bands. Estimates of the migration speed of C. inflexa were between 2 and 7 cm s⁻¹ upwards and between 4 and 7 cm s⁻¹ downwards. M. norvegica was estimated to migrate at speeds between 7 and 8 cm s⁻¹ upwards and over 11 cm s⁻¹ downwards. The consistently lower migration speeds estimated from Doppler measurements as compared with estimates obtained from measuring trajectories of back-scattering bands over time was believed to result from a methodological artefact. The Doppler measurements were nevertheless useful in a relative sense in revealing the relative speed of individuals within swarms. It was shown that individuals at the front of the upwardly migrating band of M. norvegica moved more slowly than those at the rear. These results illustrate the extra biological information that can be obtained by ADCPs compared with conventional echo-sounders.     

Role of colloidal material in the removal of 234Th in the Canada basin of the Arctic Ocean

The phase partitioning of ²³⁴Th between dissolved (<10-kiloDalton, kD), colloidal (10 kD—0.4 μm), and particulate (⩾0.5 μm) matter across a horizontal transect, from a coastal station to the deep Canada Basin, and a vertical profile in the deep Canada Basin of the western Arctic Ocean was investigated. Concentrations of suspended particulate matter (SPM), dissolved, colloidal and particulate organic carbon, particulate organic nitrogen and nutrients (silicate, phosphate and nitrate) were also measured to assess transport and scavenging processes.Total ²³⁴Th (colloidal+particulate+dissolved) indicated deficiencies relative to secular equilibrium with its parent, ²³⁸U in the upper 100 m, which suggests active scavenging of ²³⁴Th onto particle surfaces. In contrast, at depths >200 m, general equilibrium existed between total ²³⁴Th and ²³⁸U. The inventory of SPM and the specific activity of particulate ²³⁴Th in the Canada Basin was about an order of magnitude higher than the profile reported for the Alpha Ridge ice camp station. This higher concentration of SPM in the southwestern Canada Basin is likely derived from ice-rafted sedimentary particles. Inventories of nutrients, and dissolved organic carbon and nitrogen in the upper 100 m of the Canada Basin are comparable to the other estimates for the central Arctic Ocean. Comparison of the mass concentrations of colloidal and filter-retained particulate matter as well as the activity of ²³⁴Th in these phases indicates that only a very small component of the colloidal material is actively involved in Th scavenging. Lower values of the conditional partition coefficient between the colloidal and dissolved phase indicate that the Arctic colloids are less reactive than colloidal material from other regions. The conditional partition coefficient between the filter-retained and dissolved phases (K_f) is generally higher than that for other regions, which is attributed to the higher complexation capacity of glacio-marine sedimentary particles in these waters. The ²³⁴Th-derived export of POC for the shelf and deep Canada Basin ranges between 5.6 and 6.5 mmol m⁻² d⁻¹, and is in agreement with other estimates reported for the central Arctic Ocean and Beaufort Sea.