Surface Waters of the NW Iberian Margin: Upwelling on the Shelf versus Outwelling of Upwelled Waters from the Rías Baixas

A set of hydrographic surveys were carried out in the Ría of Vigo (NW Spain) at 2–4 d intervals during four 2–3 week periods in 1997, covering contrasting seasons. Residual exchange fluxes with the adjacent shelf were estimated with a 2-D, non-steady-state, salinity–temperature weighted box model. Exchange fluxes consist of a steady-state term (dependent on the variability of continental runoff) and a non-steady-state term (dependent on the time changes of density gradients in the embayment). More than 95% of the short-time-scale variability of the exchange fluxes in the middle and outer ría can be explained by the non-steady-state term that, in turns, is correlated (R²>75%) with the offshore Ekman transport. Conversely, 96% of the variability of exchange fluxes in the inner ría rely on the steady-state term. The outer and middle ría are under the direct influence of coastal upwelling, which enhances the positive residual circulation pattern by an order of magnitude: from 10²to 10³ m³s⁻¹. On the contrary, downwelling provokes a reversal of the circulation in the outer ría. The position of the downwelling front along the embayment depends on the relative importance of Ekman transport (Q_x, m³s⁻¹km⁻¹) and continental runoff (R, m³s⁻¹). When Q_x/ R>7±2 the reversal of the circulation affects the middle ría. Our results are representative for the ‘Rías Baixas’, four large coastal indentations in NW Spain. During the upwelling season (spring and summer), 60% of shelf surface waters off the ‘Rías Baixas’ consist of fresh Eastern North Atlantic Central Water (ENACW) upwelled in situ. The remaining 40% consists of upwelled ENACW that previously enters the rías and it is subsequently outwelled after thermohaline modification. During the downwelling season (autumn and winter), 40% of the warm and salty oceanic subtropic surface water, which piled on the shelf by the predominant southerly winds, enters the rías.     

Influence of upwelling and river runoff interaction on phytoplankton assemblages in a Middle Galician Ria and Comparison with northern and southern rias (NW Iberian Peninsula)

The first oceanographic research (hydrography, nutrient salts, chlorophyll, primary production and phytoplankton assemblages) in a Middle Galician Ria was carried out in Corme-Laxe during 2001, just a year before the Prestige oil spill, being the only reference to evaluate eventual changes in the phytoplankton community. Due to the small size of this ria (6.5 km²), oceanographic processes were driven by the continental water supplied by Anllons River during the wet season (20–30 m³ s⁻¹ in winter), and the strong oceanic influence from the nearby shelf during the dry season. The annual cycle showed a spring bloom with high levels of chlorophyll (up to 14 μg Chl-a L⁻¹) and primary production (3 g C m⁻² d⁻¹) and a summer upwelling bloom (up to 8 μg Chl-a L⁻¹ and 10 g C m⁻² d⁻¹) where the proximity of the Galician upwelling core (<13.5 °C at sea surface) favors the input of upwelled seawater (up to 9 μM of nitrate and silicate) to the bottom ria layer, even during summer stratification events (primary production around 2 g C m⁻² d⁻¹). Thus, phytoplankton assemblages form a “continuum” from spring to autumn with a predominance of diatoms and overlapping species between consecutive periods; only in autumn dinoflagellates and flagellates characterized the phytoplankton community. In the Middle Rias as Corme-Laxe, the nutrient values, Chl-a, primary production and phytoplankton abundance for productive periods were higher than those reported for the Northern (Ria of A Coruña) and Southern Rias (Ria of Arousa) for year 2001; this suggests the importance of the hydrographic events occurring in the zone of maximum upwelling intensity of the Western Iberian Shelf, where a lack of annual cycles studies exists.     

Short-term variability of fCO2 in seawater and air–sea CO2 fluxes in a coastal upwelling system (Ría de Vigo, NW Spain)

Coastal upwelling systems are regions with highly variable physical processes and very high rates of primary production and very little is known about the effect of these factors on the short-term variations of CO₂ fugacity in seawater (fCO₂^w). This paper presents the effect of short-term variability (<1 week) of upwelling–downwelling events on CO₂ fugacity in seawater (fCO₂^w), oxygen, temperature and salinity fields in the Ría de Vigo (a coastal upwelling ecosystem). The magnitude of fCO₂^w values is physically and biologically modulated and ranges from 285 μatm in July to 615 μatm in October. There is a sharp gradient in fCO₂^w between the inner and the outer zone of the Ría during almost all the sampling dates, with a landward increase in fCO₂^w.CO₂ fluxes calculated from local wind speed and air–sea fCO₂ differences indicate that the inner zone is a sink for atmospheric CO₂ in December only (−0.30 mmol m⁻² day⁻¹). The middle zone absorbs CO₂ in December and July (−0.05 and −0.27 mmol·m⁻² day⁻¹, respectively). The oceanic zone only emits CO₂ in October (0.36 mmol·m⁻² day⁻¹) and absorbs at the highest rate in December (−1.53 mmol·m⁻² day⁻¹).     

Nutrient and carbon removal ratios and fluxes in the Ross Sea, Antarctica

Net community production (NCP) and nutrient deficits (Def(X)) were calculated using decreases in dissolved CO₂ and nutrient concentrations due to biological removal in the upper 200 m of the water column during four cruises in the Ross Sea, Antarctica along 76°30′S in 1996 and 1997. A comparison to excess dissolved and particulate organic carbon showed close agreement between surplus total organic carbon (TOC) and NCP during bloom initiation and productivity maximum; however, when TOC values had returned to low wintertime values NCP was still significantly above zero. This seasonal NCP, 3.9±1 mol C m⁻², must be equivalent to the particle export to depths greater than 200 m over the whole productive season. We estimate that the annual export was 55±22% of the seasonal maximum in NCP. The fraction of the seasonal maximum NCP that is exported through 200 m is significantly higher than that measured by moored sediment traps at a depth of 206 m. The removal of carbon, nitrate and phosphate (based on nutrient disappearance since early spring) and their ratios showed significant differences between regions dominated by diatoms and regions dominated by the haptophyte Phaeocystis antarctica. While the ΔC/ΔN removal ratio was similar (7.8±0.2 for diatoms and 7.2±0.1 for P. antarctica), the ΔN/ΔP and ΔC/ΔP removal ratios for diatoms (10.1±0.3 and 80.5±2.3) were significantly smaller than those of P. antarctica (18.6±0.4 and 134.0±4.7). The similarity in ΔC/ΔN removal ratios of the two assemblages suggests that preferential uptake of phosphate by diatoms caused the dramatic differences in ΔC/ΔP and ΔN/ΔP removal ratios. In contrast to low ΔC/ΔP and ΔN/ΔP removal ratio in diatom-dominated areas early in the growing season, deficit N/P and C/P ratios in late autumn indicate that the elemental stochiometry of exported organic matter did not deviate significantly from traditional Redfield ratios. Changes in biologically utilized nutrient and carbon ratios over the course of the growing season indicated either a substantial remineralization of phosphate or a decrease in phosphate removal relative to carbon and total inorganic nitrogen over the bloom period. The species dependence in C/P ratios, and the relative constancy in the C/N ratios, makes N a better proxy of biological utilization of CO₂.     

Nutrient dynamics and seasonal succession of phytoplankton assemblages in a Southern European Estuary: Ria de Aveiro, Portugal

The spatial and temporal dynamics of dissolved inorganic nitrogen, dissolved phosphate, dissolved silica and chlorophyll a were measured seasonally at eight stations in the Ria de Aveiro. Between December 2000 and September 2001, the seasonal succession of phytoplankton assemblages, inferred after the spatial and seasonal variation of silica and of chlorophyll a concentrations, showed that diatoms (μmol Si L⁻¹) dominated from late autumn until early spring, while chlorophytes (μg Chl a L⁻¹) bloomed during late spring and summer. The Si:N:P ratios and Si concentrations indicated no seasonal depletion in dissolved silica, as in other temperate systems, possibly because of abnormal precipitation and flood events prolonging the supply of dissolved Si to the system. The Si:N:P ratios suggested P limitation at the system level. Despite the relative proportions of available nutrients, the mean phosphorus concentration (5.3 μmol L⁻¹) was above the reported half-saturation constants for P uptake by phytoplankton. Thus, in Ria de Aveiro, the seasonal succession of phytoplankton assemblages may also be dependent on the grazing capacity of the pelagic community through top-down regulation.     

Seasonal Variations of Nutrients, Seston and Phytoplankton, and Upwelling Intensity off La Coruña (NW Spain)

This study describes the main seasonal stages in oceanographic conditions and phytoplankton off La Coruña (Galicia, NW Spain), during 1991 and 1992, based mainly on monthly cruises near the coast. Upwelling conditions were studied using an upwelling index calculated from local winds. The Galician coast is affected by a long upwelling season for most of the year. The upwelling pulses interact with the thermal stratification-mixing cycle of surface waters, primarily affecting the dynamics of phytoplankton. In addition, the presence of water masses of different salinity in the subsurface layers changes the stratification of the water column. The less-saline North Atlantic Central Water (NACW) was normally associated with upwelling events during summer. However, on several occasions during the study, the presence of Eastern North Atlantic Water (ENAW) of subtropical origin was observed with salinities up to 36·22 and temperatures between 13 and 14 °C.Observations were grouped into five main stages related to the degree of surface stratification and characteristics of phytoplankton communities. These stages were recognized in both annual cycles, and were termed: winter mixing, spring and autumn blooms, summer upwelling, thermal stratification and special events (red tides and downwelling). A homogeneous water column was the main characteristic of the winter stage, with high nutrient concentrations and low phytoplankton biomass. Eastern North Atlantic Water appeared at the end of this stage, which lasted from November to February. The spring and autumn blooms occurred along with weak thermohaline gradients at the surface, producing high phytoplankton concentrations. Favourable upwelling conditions and the presence of ENAW in a subsurface layer were the factors that most likely induced earlier blooms, while thermal gradients developed at the surface could have been more important for later blooms. Upwelling events during summer were related to a reduction in the depth of the surface mixed layer as the pycnocline moved upwards, and can produce significant phytoplankton accumulations. These summer blooms interrupted the thermal stratification stage, characterized by low nutrient and phytoplankton concentrations at the surface. The dominant phytoplankton in the study was composed mainly of diatoms, especially during blooms. However, a proliferation of red-tide dinoflagellates was observed along with weak upwelling conditions in late summer. Also in late summer, strong downwelling conditions caused the accumulation of warmer shelf waters inshore, inducing the sinking of particulate matter produced at the surface.     

Seasonal evolution of the transverse thermohaline asymmetry in the Ria de Pontevedra (northwestern Spain)

A quasi-permanent transverse asymmetry in the thermohaline properties of the Ria de Pontevedra (northwestern Spain) was measured from October 1997 to October 1998 during 23 cruises. On average, the density increased form north to south. This asymmetry was maintained by salinity during the wet season, and by salinity and temperature during the dry season. The density difference between coasts (Δσ_t = σ_tsouth − σ_tnorth) and the river discharge were positively correlated (correlation coefficient, approximately 0.6) throughout the year. The density difference was negatively correlated with the upwelling index during the wet season (correlation coefficient, approximately −0.45) and positively correlated during the dry season (correlation coefficient, approximately 0.55).     

C : N ratios in the mixed layer during the productive season in the northeast Atlantic Ocean

Redfield stoichiometry has proved a robust paradigm for the understanding of biological production and export in the ocean on a long-term and a large-scale basis. However, deviations of carbon and nitrogen uptake ratios from the Redfield ratio have been reported. A comprehensive data set including all carbon and nitrogen pools relevant to biological production in the surface ocean (DIC, DIN, DOC, DON, POC, PON) was used to calculate seasonal new production based on carbon and nitrogen uptake in summer along 20°W in the northeast Atlantic Ocean. The 20°W transect between 30 and 60°N covers different trophic states and seasonal stages of the productive surface layer, including early bloom, bloom, post-bloom and non-bloom situations. The spatial pattern has elements of a seasonal progression. We also calculated exported production, i.e., that part of seasonal new production not accumulated in particulate and dissolved pools, again separately for carbon and nitrogen. The pairs of estimates of `seasonal new production’ and `exported production’ allowed us to calculate the C : N ratios of these quantities. While suspended particulate matter in the mixed layer largely conforms to Redfield stoichiometry, marked deviations were observed in carbon and nitrogen uptake and export with progressing season or nutrient depletion. The spring system was characterized by nitrogen overconsumption and the oligotrophic summer system by a marked carbon overconsumption. The C : N ratios of seasonal new as well as exported production increase from early bloom values of 5–6 to values of 10–16 in the post-bloom/oligotrophic system. The summertime accumulation of nitrogen-poor dissolved organic matter can explain only part of this shift.     

Effects of upwelling,tides and biological processes on the inorganic carbon system of a coastal lagoon in Baja California

The role of coastal lagoons and estuaries as sources or sinks of inorganic carbon in upwelling areas has not been fully understood. During the months of May–July, 2005, we studied the dissolved inorganic carbon system in a coastal lagoon of northwestern Mexico during the strongest period of upwelling events. Along the bay, different scenarios were observed for the distributions of pH, dissolved inorganic carbon (DIC) and apparent oxygen utilization (AOU) as a result of different combinations of upwelling intensity and tidal amplitude. DIC concentrations in the outer part of the bay were controlled by mixing processes. At the inner part of the bay DIC was as low as 1800 μmol kg⁻¹, most likely due to high water residence times and seagrass CO₂ uptake. It is estimated that 85% of San Quintín Bay, at the oceanic end, acted as a source of CO₂ to the atmosphere due to the inflow of CO₂-rich upwelled waters from the neighboring ocean with high positive fluxes higher than 30 mmol C m⁻² d⁻¹. In contrast, there was a net uptake of CO₂ and HCO₃⁻ by the seagrass bed Zostera marina in the inner part of the bay, so the pCO₂ in this zone was below the equilibrium value and slightly negative CO₂ fluxes of −6 mmol C m⁻² d⁻¹. Our positive NEP and ΔDIC values indicate that Bahía San Quintín was a net autotrophic system during the upwelling season during 2005.     

Grazing impact of mesozooplankton in an upwelling region off northern California, 2000–2003

Mesozooplankton (>200 μm) grazing impact (% phytoplankton standing crop consumed d⁻¹) was determined by the gut fluorescence method during three springs (2000, 2001 and 2002) and two winters (2002 and 2003) in a coastal upwelling region off northern California. Wind events, in terms of both magnitude and duration, varied inter-annually and seasonally and included both upwelling-favorable and relaxation events. Grazing impact of mesozooplankton also varied inter-annually and seasonally, and was highest during June 2000 (mean=129% of standing crop d⁻¹), a prolonged period of wind “relaxation” and phytoplankton bloom. In contrast, mean grazing impact was lower during periods of stronger, more persistent winds, more active upwelling, greater cross-shelf transport, and lower chlorophyll concentration (25% and 38% in May–June 2001 and 2002, respectively). Wintertime conditions (January 2002 and 2003) were characterized by weakly upwelling or downwelling-favorable winds, low chlorophyll concentration, and lower mean mesozooplankton grazing impact (13% and 12%, respectively). The larger (>500 μm) size class contributed proportionally more to total mesozooplankton (>200 μm) grazing impact than the smaller (200–500 μm) size class during all sampling periods except spring 2002. These results suggest that mesozooplankton grazing impact is higher in spring than in winter, and that during the spring upwelling season, grazing is higher during periods of wind relaxation (weak upwelling) than during periods of stronger upwelling. Further, these results suggest an important role of mesozooplankton grazers on phytoplankton dynamics in the upwelling region off northern California.     

Remineralization and accumulation of organic carbon and nitrogen in marine sediments of eutrophic bays: the case of the Bay of Concepcion, Chile

The Bay of Concepcion (36°40′S; 73°02′W) is a semi-enclosed and shallow embayment in which biogeochemical processes are seasonally coupled to coastal upwelling during the austral spring and summer. The nutrient cycle in the bay is complex due to the combined effects of a pronounced O₂ minimum layer and high nutrient concentrations both originating from subsurface equatorial water during coastal upwelling and a rapid rate of sediment nutrient recycling. The sediments are characterized by a high content of organic matter mainly due to the extremely high rates of phytoplankton production and deposition. During the upwelling period, a black flocculent layer frequently covers the sediment–water interface in the inner part of the bay where an extensive mat of Beggiatoa spp. develops. Three approaches are used to analyse the extent to which the benthic system recycles or retains nutrients at two stations, located at the centre (station C, St. C) and mouth (station B, St. B) of the bay for a 1-year period (March 1996–1997): (1) estimation of C and N remineralization rates based on SO₄²⁻ reduction measurements, (2) calculation of C and N turnover rates using a diagenetic model applied to total organic carbon and total nitrogen vertical distributions and, (3) construction of C and N budgets from direct measurements of sedimentation (from a sediment trap) and estimates of the C and N burial rates. Depth-integrated SO₄²⁻ reduction rates varied between 3.4 (winter) and 25.5 (summer) mmol m⁻² d⁻¹. Estimated C and N oxidation rates ranged between 7.9 and 87.8 mol C m⁻² yr⁻¹ and between 0.9 and 6.9 mol N m⁻² yr⁻¹, respectively. Each approach yielded minor differences in the C and N remineralization rates (and also minor differences between both studied stations), except when the kinetic model was applied to C and N distribution without including the presence of the flocculent layer. The rates of carbon oxidation and sulphate reduction were considerably higher than in other coastal sediments with similar depositional regime. The C and N burial rates were 2.23 and 0.21 (St. C) and 1.30 and 0.09 (St. B) mol m⁻² yr⁻¹, respectively. The C/N ratio of the buried fraction was ca. 10.6 at St. C and 14.4 at St. B. Because the observed differences in burial rates could not be ascribed to distinctive depositional (both stations have similar sediment accumulation rates) and oceanographic (similar O₂ concentration and hydrography) conditions, differences may be due to in part spatial heterogeneity in the supply of organic matter. The degree of preservation of organic matter as plankton detritus and nitrogen accumulating bacterial biomass associated with Beggiatoa spp. at St. C may also be involved.     

Primary production and nutrient assimilation in the Iberian upwelling in August 1998

Primary production was measured during two Lagrangian experiments in the Iberian upwelling. The first experiment, in a body of upwelled water, measured day-to-day changes in phytoplankton activity as the water mass moved south along the shelf break. Nutrient concentrations decreased over a five day period, with concomitant increases in phytoplankton biomass. Initially the maximum phytoplankton biomass was in the upper 10m but after four days, a sub-surface chlorophyll maximum was present at 30m. Depth-integrated primary production at the beginning of the experiment was 70mmolC.m⁻².d⁻¹ (838mgC.m⁻².d⁻¹) and reached a maximum of 88mmolC.m⁻².d⁻¹ (1053mgC.m⁻².d⁻¹) on day 3. On day 1, the picoplankton fraction (<2μm) was slightly more productive than larger (>5μm) phytoplankton, but the increase in overall production during the drift experiment was by these larger cells. Nitrate was the dominant nitrogen source. As nutrient concentrations declined, ammonium became increasingly more important as a nitrogen source and the f-ratio decreased from 0.7 to 0.5. Picoplankton cells (<2μm) were responsible for most (65–80%) of the ammonium uptake. The C:N:P uptake ratios were very close to the Redfield ratio for the first four days but as nutrients became depleted high C:N uptake ratios (11 to 43) were measured. Over the period of the experiment, nitrate concentration within the upper 40m decreased by 47.91mmolN.m⁻². In vitro estimates, based on ¹⁵N nitrate uptake, accounted for 56% of the decrease in nitrate concentration observed in the drifting water mass. Ammonium uptake over the same four day period was 16.28mmolN.m⁻², giving a total nitrogen uptake of 43.18mmolN.m⁻².In the second experiment, an offshore filament was the focus and a water mass was sampled as it moved offshore. Nutrient concentrations were very low (nitrate was <10nmol l⁻¹ and ammonium was 20–40nmol l⁻¹). Primary production rate varied between 36mmolC.m⁻².d⁻¹ (436mgC.m⁻².d⁻¹) and 21mmolC.m⁻².d⁻¹ (249mgC.m⁻².d⁻¹). Picophytoplankton was the most productive fraction and was responsible for a constant proportion (ca 0.65) of the total carbon fixation. Uptake rates of both nitrate and ammonium were between 10 and 20% of those measured in the upwelling region. Urea could be a very significant nitrogen source in these waters with much higher uptake rates than nitrate or ammonium; urea turnover times were ca. one day but the source of the urea remains unknown. Urea uptake had a profound effect on calculated f ratios. If only nitrate and ammonium uptake was considered, f ratios were calculated to be 0.42–0.46 but inclusion of urea uptake reduced the f ratio to <0.1. The primary production of this oligotrophic off-shore filament was driven by regenerated nitrogen.     

南海浅层经向翻转环流及相关的内部水体流动

The structure of the annual-mean shallow meridional overturning circulation(SMOC) in the South China Sea(SCS) and the related water movement are investigated,using simple ocean data assimilation(SODA) outputs.The distinct clockwise SMOC is present above 400 m in the SCS on the climatologically annual-mean scale,which consists of downwelling in the northern SCS,a southward subsurface branch supplying upwelling at around 10°N and a northward surface flow,with a strength of about 1×10~6 m~3/s.The formation mechanisms of its branches are studied separately.The zonal component of the annual-mean wind stress is predominantly westward and causes northward Ekman transport above 50 m.The annual-mean Ekman transport across 18°N is about 1.2×10~6 m~3/s.An annual-mean subduction rate is calculated by estimating the net volume flux entering the thermocline from the mixed layer in a Lagrangian framework.An annual subduction rate of about 0.66×10~6m~3/s is obtained between 17° and 20°N,of which 87% is due to vertical pumping and 13% is due to lateral induction.The subduction rate implies that the subdution contributes significantly to the downwelling branch.The pathways of traced parcels released at the base of the February mixed layer show that after subduction water moves southward to as far as 11°N within the western boundary current before returning northward.The velocity field at the base of mixed layer and a meridional velocity section in winter also confirm that the southward flow in the subsurface layer is mainly by strong western boundary currents.Significant upwelling mainly occurs off the Vietnam coast in the southern SCS.An upper bound for the annual-mean net upwelling rate between 10° and 15°N is 0.7×10~6m~3/s,of which a large portion is contributed by summer upwelling,with both the alongshore component of the southwest wind and its offshore increase causing great upwelling.     

Responses of nutrient biogeochemistry and nitrogen cycle to seasonal upwelling in coastal waters of the eastern Hainan Island

The coastal upwelling has profound influence on the surrounding ecosystem by supplying the nutrient-replete water to the euphotic zone. Nutrient biogeochemistry was investigated in coastal waters of the eastern Hainan Island in summer 2015 and autumn 2016. From perspectives of nutrient dynamics and physical transport, the nutrient fluxes entered the upper 50 m water depth（between the mixed layer and the euphotic zone） arisen from the upwelling were estimated to be 2.5-5.4 mmol/（m²·d）,...     

Marine dissolved organic matter: can its C : N ratio explain carbon overconsumption?

Carbon overconsumption, i.e. the consumption of inorganic carbon relative to inorganic nitrogen in excess of the Redfield ratio at the sea surface, was examined in relation to the dynamics of dissolved organic carbon and nitrogen (DOC and DON) in the northeast Atlantic. We observed the presence of N-poor dissolved organic matter (DOM) in surface water during summer, requiring the consumption of inorganic carbon and nitrogen in a ratio exceeding the Redfield ratio. The C : N ratio of bulk DOM is not only different from the Redfield ratio but also variable, i.e. no fixed conversion factor of C and N exists where DOM is important in C and N transformations. The existence of N-poor DOM is recognized as a feature typical of oligotrophic systems. At the same time, the C : N ratios of particles conform to Redfield stoichiometry as does deep-ocean chemistry. The implications of this finding are discussed, the conclusion being that, while DOM buildup contributes to CO₂ drawdown seasonally, its impact on long-term carbon and nitrogen balance of the ocean is small.     

Atmospherically-promoted photosynthetic activity in a well-mixed ecosystem: Significance of wet deposition events of nitrogen compounds

Wet atmospheric deposition of dissolved N, P and Si species is studied in well-mixed coastal ecosystem to evaluate its potential to stimulate photosynthetic activities in nutrient-depleted conditions. Our results show that, during spring, seawater is greatly depleted in major nutrients: Dissolved Inorganic Nitrogen (DIN), Dissolved Inorganic Phosphorus (DIP) and Silicic acid (Si), in parallel with an increase of phytoplanktonic biomass. In spring (March–May) and summer (June–September), wet atmospheric deposition is the predominant source (>60%, relative to riverine contribution) for nitrates and ammonium inputs to this N-limited coastal ecosystem. During winter (October–February), riverine inputs of DIN predominate (>80%) and are annually the most important source of DIP (>90%). This situation allows us to calculate the possibility for a significant contribution to primary production in May 2003, from atmospheric deposition (total input for DIN ≈300 kg km⁻² month⁻¹). Based on usual Redfield ratios and assuming that all of the atmospheric-derived N (AD-N) in rainwater is bioavailable for phytoplankton growth, we can estimate new production due to AD-N of 950 mg C m⁻² month⁻¹, during this period of depletion in the water column. During the same episode (May 2003), photosynthetic activity rate, considered as gross primary production, was estimated to approximately 30 300 mg C m⁻² month⁻¹. Calculation indicates that new photosynthetic activity due to wet atmospheric inputs of nitrogen could be up to 3%.     

The effect of water transport on nitrogen flow through a kelp-bed community

A simulation model is used to investigate possible ecological effects of up- and downwelling on nitrogen flows through a kelp-bed food web off the Cape Peninsula, South Africa. The model depicts the flow of nitrogen, which is often limiting in marine ecosystems, from kelps, other macrophytes and phytoplankton, through filter-feeders to carnivores, with a feedback loop via faeces and bacteria to detritus and filter-feeders. When modelled as a closed system, bacteria associated with detritus and animal faeces form a large component of the particulate nitrogen available to filter-feeders, and the faeces feedback loop dominates nitrogen flow. When measured rates of water transport are incorporated into the model, bacteria have little opportunity to accumulate before being transported out of the system. Animal faeces and kelp detritus are the dominant filter-feeder food components under upwelling conditions, whereas phytoplankton is the major contributor to particulate organic nitrogen under downwelling conditions. When realistic pulses of upwelling/downwelling derived from wind indices are used as model input, filter-feeders are shown to decline during the summer upwelling season when much potential food is advected out of the system, and they increase during the winter when downwelling conditions are more prevalent, bringing in nitrogen-rich phytoplankton from the blooms developing offshore.     

Seasonal dynamics of inorganic nutrients and phytoplankton biomass in the NW Alboran Sea

The seasonal dynamics of inorganic nutrients and phytoplankton biomass (chlorophyll a), and its relation with hydrological features, was studied in the NW Alboran Sea during four cruises conducted in February, April, July and October 2002. In the upper layers, the seasonal pattern of nutrient concentrations and their molar ratios (N:Si:P) was greatly influenced by hydrological conditions. The higher nutrient concentrations were observed during the spring cruise (2.54 μM NO₃⁻, 0.21 μM PO₄³⁻ and 1.55 μM Si(OH)₄, on average), coinciding with the increase of salinity due to upwelling induced by westerlies. The lowest nutrient concentrations were observed during summer (<0.54 μM NO₃⁻, 0.13 μM PO₄³⁻ and 0.75 μM Si(OH)₄, on average), when the lower salinities were detected. Nutrient molar ratios (N:Si:P) followed the same seasonal pattern as nutrient distribution. During all the cruises, the ratio N:P in the top 20 m was lower than 16:1, indicating a NO₃⁻ deficiency relative to PO₄³⁻. The N:P ratio increased with depth, reaching values higher than 16:1 in the deeper layers (200–300 m). The N:Si ratio in the top 20 m was lower than 1:1, excepting during spring when N:Si ratios higher than 1:1 were observed in some stations due to the upwelling event. The N:Si ratio increased with depth, showing a maximum at 50–100 m (>1.5:1), which indicates a shift towards Si-deficiency in these layers. The Si:P ratio was much lower than 16:1 throughout the water column during the four cruises. In general, the spatial and seasonal variation of phytoplankton biomass showed a strong coupling with hydrological and chemical fields. The higher chlorophyll a concentrations at the depth of the chlorophyll maximum were found in April (2.57 mg m⁻³ on average), while the lowest phytoplankton biomass corresponded to the winter cruise (0.74 mg m⁻³ on average). The low nitrate concentrations together with the low N:P ratios found in the upper layers (top 20 m) during the winter, summer and autumn cruises suggest that N-limitation could occur in these layers during great part of the year. However, N-limitation during the spring cruise was temporally overcome by nutrient enrichment caused by an intense wind-driven upwelling event.     

Summer algal blooms in a coastal ecosystem: the role of atmospheric deposition versus entrainment fluxes

The nitrogen inputs from atmospheric deposition and bottom water entrainment to the surface layer were modelled in the summer period (May–September) over a 11-year period (1989–1999) and compared to investigate the significance of these fluxes for generating blooms in the Kattegat. In the summer periods the average atmospheric deposition was 2.81 mg N m⁻² d⁻¹ compared to average entrainment fluxes of 5.42 mg N m⁻² d⁻¹, 1.21 mg N m⁻² d⁻¹ and 1.15 mg N m⁻² d⁻¹ for the northern, central and southern part of the Kattegat, respectively. Atmospheric nitrogen deposition alone could not sustain biomass increases associated with observed blooms and entrainment fluxes dominated the high nitrogen inputs to the surface layer. The potential for a bloom through growth was typically obtained after several days of high nitrogen inputs from entrainment in the frontal area of the northern Kattegat and to some extent from atmospheric deposition. The modelled nitrogen input in this area could account directly for 30% of the observed blooms in the Northern sub-basin, and through advective transport 24% and 19% of the observed blooms in the central and southern Kattegat. The direct nitrogen inputs through atmospheric deposition and entrainment to the central and southern sub-basins were small and could not be linked to any bloom observation.     

A nitrate and silicate budget in the equatorial Pacific Ocean: a coupled physical–biological model study

A coupled physical–biological model was developed to simulate the low-silicate, high-nitrate, and low-chlorophyll (LSHNLC) conditions in the equatorial Pacific Ocean and used to compute a detailed budget in the Wyrtki box (5°N–5°S, 180–90°W) for the major sources and cycling of nitrogen and silicon in the equatorial Pacific. With the incorporation of biogenic silicon dissolution, NH₄ regeneration from organic nitrogen and nitrification of ammonia in the model, we show that silicon recycling in the upper ocean is less efficient than nitrogen. As the major source of nutrients to the equatorial Pacific, the Equatorial Undercurrent provides slightly less Si(OH)₄ than NO₃ to the upwelling zone, which is defined as 2.5°N–2.5°S. As a result, the equatorial upwelling supplies less Si(OH)₄ than NO₃ into the euphotic zone in the Wyrtki box, having a Si/N supply ratio of about 0.85 (2.5 vs. 2.96 mmolm⁻² day⁻¹). More Si(OH)₄ than NO₃ is taken up with a Si/N ratio of 1.17 (2.72 vs. 2.33 mmolm⁻² day⁻¹) within the euphotic zone. The difference between upwelling supply and biological uptake is balanced by nutrient regeneration and horizontal advection. Excluding regeneration, the net silicate and nitrate uptakes are nearly equal (1.76 vs. 1.84 mmolm⁻² day⁻¹). However, biogenic silica export production is slightly higher than organic nitrogen (1.74 vs. 1.59 mmolm⁻² day⁻¹) following a 1.1 Si/N ratio. In the central equatorial Pacific, low silicate concentrations limit diatom growth; therefore non-diatom new production accounts for most of the new production. Higher silicate supply in the east maintains elevated diatom growth rates and new production associated with diatoms dominate upwelling zone. In contrast, the new production associated with small phytoplankton is nearly constant or decreases eastward along the equator. The total new production has a higher rate in the east than in the west, following the pattern of surface silicate. This suggests that silicate regulates the diatom production, total new production, and thereby carbon cycle in this area. The modeled mean primary production is 48.4 mmolCm⁻² day⁻¹, representing the lower end of direct field measurements, while new production is 15.0 mmolCm⁻² day⁻¹, which compares well with previous estimates.