WEST: A northern California study of the role of wind-driven transport in the productivity of coastal plankton communities

The “Wind Events and Shelf Transport” (WEST) program was an interdisciplinary study of coastal upwelling off northern California in 2000–03. WEST was comprised of modeling and field observations. The primary goal of WEST was to better describe and understand the competing influences of wind forcing on planktonic productivity in coastal waters. While increased upwelling-favorable winds lead to increased nutrient supply, they also result in reduced light exposure due to deeper surface mixed layers and increased advective loss of plankton from coastal waters. The key to understanding high levels of productivity, amidst these competing responses to wind forcing, is the temporal and spatial structure of upwelling. Temporal fluctuations and spatial patterns allow strong upwelling that favors nutrient delivery to be juxtaposed with less energetic conditions that favor stratification and plankton blooms. Observations of winds, ocean circulation, nutrients, phytoplankton and zooplankton off Bodega Bay and Point Reyes (38°N) were combined with model studies of winds, circulation and productivity. This overview of the WEST program provides an introduction to the WEST special issue of Deep-Sea Research, including the motivation for WEST, a summary of study components, an integrative synthesis of major research results to-date, and background on conditions during field studies in May–June 2001 (the upwelling period on which this special issue is focused).     

Nutrient contributions to the Santa Barbara Channel, California, from the ephemeral Santa Clara River

The Santa Clara River delivers nutrient rich runoff to the eastern Santa Barbara Channel during brief (1–3 day) episodic events. Using both river and oceanographic measurements, we evaluate river loading and dispersal of dissolved macronutrients (silicate, inorganic N and P) and comment on the biological implications of these nutrient contributions. Both river and ocean observations suggest that river nutrient concentrations are inversely related to river flow rates. Land use is suggested to influence these concentrations, since runoff from a subwatershed with substantial agriculture and urban areas had much higher nitrate than runoff from a wooded subwatershed. During runoff events, river nutrients were observed to conservatively mix into the buoyant, surface plume immediately seaward of the Santa Clara River mouth. Dispersal of these river nutrients extended 10s of km into the channel. Growth of phytoplankton and nutrient uptake was low during our observations (1–3 days following runoff), presumably due to the very low light levels resulting from high turbidity. However, nutrient quality of runoff (Si:N:P = 16:5:1) was found to be significantly different than upwelling inputs (13:10:1), which may influence different algal responses once sediments settle. Evaluation of total river nitrate loads suggests that most of the annual river nutrient fluxes to the ocean occur during the brief winter flooding events. Wet winters (such as El Niño) contribute nutrients at rates approximately an order-of-magnitude greater than “average” winters. Although total river nitrate delivery is considerably less than that supplied by upwelling, the timing and location of these types of events are very different, with river discharge (upwelling) occurring predominantly in the winter (summer) and in the eastern (western) channel.     

Nearshore retention of upwelled waters north and south of Point Reyes (northern California)—Patterns of surface temperature and chlorophyll observed in CoOP WEST

Retentive embayments can be found near capes in upwelling regions, where they stand out as relatively warm features with higher chlorophyll a concentrations than surrounding waters. Within the area of the “Wind Events and Shelf Transport” study site (WEST) from the Gulf of the Farallones to Point Arena (37.5–39°N and 122.5–124°W), we describe two retentive embayments, extending approximately 20 km north of Point Reyes, and 30 km south of Point Reyes in the northern Gulf of the Farallones. These areas are identifiable from chlorophyll a (SeaWiFS) and sea-surface temperature (MODIS) satellite radiometry. Additional data from moorings and drifters were used to further characterize these retentive features. The persistence of these features is on time scales between 2 and 10 days, determined from time-series analysis of mooring data for chlorophyll and temperature. The alongshore wind stress is negatively correlated with chlorophyll and temperature mooring values. The time scale of persistence of these retentive features was similar to both the upwelling-relaxation timescale and the timescale of phytoplankton bloom development. The WEST region is notable in that the spring and summer is subject to the strongest wind stress along the US west coast, yet the coastal waters are highly productive. Based on our observations, we suggest that the apparent persistence of high biomass coastal waters in this strongly advective and wind-dominated system may be partially explained by the presence of retentive features identified in this study.     

Effects of variable winds on biological productivity on continental shelves in coastal upwelling systems

The production and distribution of biological material in wind-driven coastal upwelling systems are of global importance, yet they remain poorly understood. Production is frequently presumed to be proportional to upwelling rate, yet high winds can lead to advective losses from continental shelves, where many species at higher trophic levels reside. An idealized mixed-layer conveyor (MLC) model of biological production from constant upwelling winds demonstrated previously that the amount of new production available to shelf species increased with upwelling at low winds, but declined at high winds [Botsford, L.W., Lawrence, C.A., Dever, E.P., Hastings, A., Largier, J., 2003. Wind strength and biological productivity in upwelling systems: an idealized study. Fisheries Oceanography 12, 245–259]. Here we analyze the response of this model to time-varying winds for parameter values and observed winds from the Wind Events and Shelf Transport (WEST) study region. We compare this response to the conventional view that the results of upwelling are proportional to upwelled volume. Most new production per volume upwelled available to shelf species occurs following rapid increases in shelf transit time due to decreases in wind (i.e. relaxations). However, on synoptic, event time-scales shelf production is positively correlated with upwelling rate. This is primarily due to the effect of synchronous periods of low values in these time series, paradoxically due to wind relaxations. On inter-annual time-scales, computing model production from wind forcing from 20 previous years shows that these synchronous periods of low values have little effect on correlations between upwelling and production. Comparison of model production from 20 years of wind data over a range of shelf widths shows that upwelling rate will predict biological production well only in locations where cross-shelf transit times are greater than the time required for phytoplankton or zooplankton production. For stronger mean winds (narrower shelves), annual production falls below the peak of constant wind prediction [Botsford et al., 2003. Wind strength and biological productivity in upwelling systems: an idealized study. Fisheries Oceanography 12, 245–259], then as winds increase further (shelves become narrower) production does not decline as steeply as the constant wind prediction.     

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.     

The phytoplankton bloom response to wind events and upwelled nutrients during the CoOP WEST study

In the coastal waters off northern California, seasonal wind-driven upwelling supplies abundant nutrients to be processed by phytoplankton productivity. As part of the Coastal Ocean Processes: Wind Events and Shelf Transport (CoOP WEST) study, nutrients, CO₂, size-fractionated chlorophyll, and phytoplankton community structure were measured in the upwelling region off Bodega Bay, CA, during May–June 2000, 2001 and 2002. The ability of this ecosystem to assimilate nitrate (NO₃) and silicic acid/silicate (Si(OH)₄) and accumulate particulate material (i.e. phytoplankton) was realized in all 3 years, following short events of upwelling-favorable winds, followed by periods of relaxed winds. This was observed as phytoplankton blooms, dominated by chlorophyll in cells greater than 5 μm in diameter, that reduced the ambient nutrients to zero. These communities were located over the near-shore shelf (<100 m depth) and were dominated by diatoms. An optimal window of 3–7 days of relaxed winds, following an upwelling pulse, was required for chlorophyll accumulation. The large-celled phytoplankton that result are likely important players in coastal new production and carbon cycling.     

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.     

Coupling of biological and physical regimes across the Antarctic Polar Front as reflected by nitrogen production and recycling

Oceanographic samples were collected across the Antarctic Polar Front (APF) region in the vicinity of 60°S, 170°W during the US JGOFS program from December 1997 to March 1998. This paper reports the uptake rates of new (nitrate) and regenerated (ammonium and urea) nitrogen measured by ¹⁵N tracer techniques together with the levels of ammonium, urea-N and dissolved free amino acids (DFAAs) during December and mid-February–March. The APF was an important biological boundary, and in December rates of new (nitrate) uptake were greatest south of the APF, exceeding 10 mmol m⁻² d⁻¹ near the retreating ice edge. In February, nitrate uptake rates were an order of magnitude lower. Rates of ammonium uptake in both periods were greater in the warmer water north of the front. Nitrogen f-ratios varied from 0.50 to less than 0.05, with larger values associated with the >5 μm size fraction at the ice edge and generally lower values north of the APF. Urea was an important nitrogen source north of the APF, and lowered f-ratios there by 22% on average when included as part of total nitrogen uptake. Urea uptake was less important south of the APF. Ammonium concentrations increased dramatically south of the APF later in the season, suggesting a system dominated by regeneration. Seasonal changes in the concentrations of regenerated organic compounds such as urea and DFAAs were less obvious, although DFAAs exhibited consistent maxima in the high flow regions of the APF. A mass balance based of ammonium fluxes suggests that nitrification was significant at locations south of the APF in February. In these nitrate-replete waters, light/mixing conditions in the surface water (the Sverdrup criterion) accounted for over 50% of the variance in nitrate uptake rates. The stability responsible for higher new production south of the APF is due both to the separation of this region from the maximum zonal wind field to the north as well as to melt-water contribution from the retreating ice field. Estimated new production and exportable carbon production exceeded 500 mmol nitrate m⁻² yr⁻¹ and 40 g C m⁻² yr⁻¹, respectively, south of the APF. Thus, new production in the marginal ice zone of the Southern Ocean rivals that in coastal systems and indicates that this is an important region for export production.     

Wintertime rates of net primary production and nitrate and ammonium uptake in the southern Benguela upwelling system

The elevated levels of primary productivity associated with eastern boundary currents are driven by nutrient- rich waters upwelled from depth, such that these regions are typically characterised by high rates of nitrate-fuelled phytoplankton growth. Production studies from the southern Benguela upwelling system (SBUS) tend to be biased towards the summer upwelling season, yet winter data are required to compute annual budgets and understand seasonal variability. Net primary production (NPP) and nitrate and ammonium uptake were measured concurrently at six stations in the SBUS in early winter. While euphotic zone NPP was highest at the stations nearest to the coast and declined with distance from the shore, a greater proportion was potentially exportable from open-ocean surface waters, as indicated by the higher specific nitrate uptake rates and f-ratios (ratio of nitrate uptake to total nitrogen consumption) at the stations located off the continental shelf. Near the coast, phytoplankton growth was predominantly supported by ammonium despite the high ambient nitrate concentrations. Along with ammonium concentrations as high as 3.6 µmol l^–1, this strongly suggests that nitrate uptake in the inshore SBUS, and by extension carbon drawdown, is inhibited by ammonium, at least in winter, although this has also been hypothesised for the summer.     

Surface boundary-layer variability off Northern California, USA, during upwelling

A five-element mooring array is used to study surface boundary-layer transport over the Northern California shelf from May to August 2001. In this region, upwelling favorable winds increase in strength offshore, leading to a strong positive wind stress curl. We examine the cross-shelf variation in surface Ekman transport calculated from the wind stress and the actual surface boundary-layer transport estimated from oceanic observations. The two quantities are highly correlated with a regression slope near one. Both the Ekman transport and surface boundary layer transport imply curl-driven upwelling rates of about 3×10⁻⁴ m s⁻¹ between the 40 and 90 m isobaths (1.5 and 11.0 km from the coast, respectively) and curl-driven upwelling rates about 1.5×10⁻⁴m s⁻¹ between the 90 and 130 m isobaths (11.0 and 28.4 km from the coast, respectively). Thus curl-driven upwelling extends to at least 25 km from the coast. In contrast, upwelling driven by the adjustment to the coastal boundary condition occurs primarily inshore of the 40-m isobath. The upwelling rates implied by the differentiating the 40-m transport observations with the coastal boundary condition are up to 8×10⁻⁴ m s⁻¹. The estimated upwelling rates and the temperature–nitrate relationship imply curl-driven vertical nitrate flux divergences are about half of those driven by coastal boundary upwelling.     

Phosphorus input by upwelling in the eastern Gotland Basin (Baltic Sea) in summer and its effects on filamentous cyanobacteria

In July 2007, phosphorus input by an upwelling event along the east coast of Gotland Island and the response of filamentous cyanobacteria were studied to determine whether introduced phosphorus can intensify cyanobacterial bloom formation in the eastern Gotland Basin. Surface temperature, nutrient concentrations, phytoplankton biomass and its stoichiometry, as well as phosphate uptake rates were determined in two transects between the coasts of Gotland and Latvia and in a short grid offshore of Gotland. In the upwelling area, surface temperatures of 11–12 °C and average dissolved inorganic phosphorus (DIP) concentrations of 0.26 μM were measured. Outside the upwelling, surface temperatures were higher (15.5–16.6 °C) and DIP supplies in the upper 10 m layer were exhausted. Nitrite and nitrate concentrations (0.01–0.22 μM) were very low within and outside the upwelling region. Abundances of filamentous cyanobacteria were highly reduced in the upwelling area, accounting for only 1.4–6.0% of the total phytoplankton biomass, in contrast to 18–20% outside the upwelling. The C:P ratio of filamentous cyanobacteria varied between 32.8 and 310 in the upwelling region, most likely due to the introduction of phosphorus-depleted organisms into the upwelling water. These organisms accumulate DIP in upwelling water and have lower C:P ratios as long as they remain in DIP-rich water. Thus, diazotrophic cyanobacteria benefit from phosphorus input directly in the upwelling region. Outside the upwelling region, the C:P ratios of filamentous cyanobacteria varied widely, between 240 and 463, whereas those of particulate material in the water ranged only between 96 and 224. To reduce their C:P ratio from 300 to 35, cyanobacteria in the upwelling region had to take up 0.05 mmol m⁻³ DIP, which is about 20% of the available DIP. Thus, a larger biomass of filamentous cyanobacteria may be able to benefit from a given DIP input. As determined from the DIP uptake rates measured in upwelling cells, the time needed to reduce the C:P ratio from 300 to 35 was too long to explain the huge bloom formations that typically occur in summer. However, phosphorus uptake rates increased significantly with increasing C:P ratios, allowing phosphorus accumulation within 4–5 days, a span of time suitable for bloom formation in July and August.     

Nutrient distributions and new production in polar regions: parallels and contrasts between the Arctic and Antarctic

Polar regions are poorly understood components of global biogeochemical cycles. This paper discusses the influences of nitrate and ammonium concentrations on nitrate uptake (and hence new production), particularly with regard to data collected within marginal ice zones in the Arctic and Antarctic. Subsurface ammonium maxima in waters over 150 m are frequently encountered in the Arctic and occasionally in the Antarctic. Such maxima result from the heterotrophic remineralization of organic matter, and because stratified environments occur more frequently in the Arctic, significant concentrations of ammonium accumulate as a result of lower diffusive losses. Causal agents (bacteria vs. Zooplankton) may also be different in the Arctic from those in the Antarctic. Elevated ammonium concentrations significantly reduce nitrate uptake, and it is suggested that this nutrient interaction may play a significant role in controlling new production, particularly in open water regions. The new production of the Southern Ocean is estimated, and it is suggested that the production and flux from the surface layer is significant relative to the world's oceans.     

Bio-optical signatures and biogeochemistry from intense upwelling and relaxation in coastal California

The NSF-sponsored Coastal Ocean Processes Wind Events and Shelf Transport (WEST) experiment investigates the interplay between wind-driven transport and shelf productivity; while eastern boundary shelves are characterized by high productivity due to upward fluxes of nutrients into the euphotic zone, wind forcing also represents negative physical and biological controls via offshore transport and deep (light-limiting) mixing of primary producers. Although this interaction has been well documented for eastern boundary systems generally and for California specifically, one of the primary goals of WEST was to characterize more fully the interplay between positive and negative effects of wind stress, which result in the consistently elevated biological productivity in these shelf regions. During 3 month-long summer cruises (2000–2002) we observed extremes in upwelling/relaxation, using both in situ instrumentation and remotely sensed data. Relationships between optical and physical properties were examined, with emphasis on biogeochemical implications. During 2000, the WEST region was optically dominated by phytoplankton and covarying constituents. During 2001 and 2002, periods of more intense upwelling favorable winds, we observed a transition to optical properties dominated by detrital and inorganic materials. In all years, the continental shelf break provided a natural boundary between optically distinct shelf and open ocean waters. During 2002, we obtained discrete trace-metal measurements of particulate iron and aluminum; we develop a bio-optical proxy for acetic-acid leachable iron from backscatter and fluorescence, and demonstrate that particulate iron is not well correlated to traditional upwelling proxies such as macronutrients, temperature, and salinity. We conclude that the shelf break between ca. 100 and 200 m water depth serves as a natural break point between coastal and oceanic water masses in this region, and that the elevated biomass and productivity associated with this eastern boundary current regime is dominated by these iron rich, shallow shelf waters.     

Tropical Atlantic variability in a coupled physical–biogeochemical ocean model

A three-dimensional ocean biogeochemical model of the tropical Atlantic Ocean was run for more than half a century (1949–2000) in order to characterize the ocean biogeochemical response to variable forcing over this period. The seasonal cycle in the equatorial upwelling zone agrees reasonably well with observations and other published simulations but underestimates phytoplankton biomass under strong upwelling conditions. Away from the equator, modelled nutrient flux and biological production are maximal in each hemisphere's winter season, and appear to be proximately forced by evaporative cooling and wind stirring rather than by Ekman upwelling. The fraction of the total variance associated with the seasonal cycle is considerably smaller for modelled biogeochemical fields than for sea-surface temperature over this long simulation, and much of the biogeochemical variance is associated with interdecadal changes. The model results suggest that the tropical Atlantic became more productive following the Pacific climate shift of 1976 and remained so until about 1989. Summer surface nitrate concentrations during the 1990s were lower than those in the 1980s. The relationship between the equatorial and off-equatorial regimes may have changed following the 1976 event, with equatorial variability dominating the basin-wide variance patterns after 1976.     

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.     

Variation of carbon and nitrogen uptake capacity in a regional upwelling area around Hachijo Island

Capacities for inorganic carbon, nitrate and ammonium uptake were measured around Hachijo Island, 300 km south of Tokyo, where local upwelling occurred. The phytoplankton population inside the upwelling area had a high capacity for nitrate uptake and a low capacity for uptake of ammonium. Nutrient concentration and phytoplankton biomass were higher in the upwelling plume than outside. On a chlorophylla basis, phytoplankton populations inside the upwelling area showed a lower capacity for carbon and nitrogen uptake than those outside the upwelling. Low temperature, relatively limited availability of light caused by extensive water mixing within the upwelling plume, and the difference in species composition of phytoplankton must be considered in explaining these lower uptake capacities.     

Wintertime nutrients in the North Atlantic—new approaches and implications for new production estimates

Observations of wintertime nutrient concentrations in surface waters are scarce in the temperate and subarctic North Atlantic Ocean. Three new methods of their estimation from spring or early summer observations are described and evaluated. The methods make use of a priori knowledge of the vertical distribution of oxygen saturation and empirical relationships between nutrient concentrations and oxygen saturation. A south–north increase in surface water winter nutrient concentration is observed. Winter nitrate concentrations range from very low levels of about 0.5 μmol dm⁻³ at 33°N to about 13.5 μmol dm⁻³ at 60°N. Previous estimates of winter nitrate concentrations have been overestimates by up to 50%. At the Biotrans Site (47°N, 20°W), a typical station in the temperate Northeast Atlantic, a mean winter nitrate concentration of 8 μmol dm⁻³ is estimated, compared to recently published values between 11 and 12.5 μmol dm⁻³. It is shown that most of the difference is due to a contribution of remineralised nitrate that had not been recognized in previous winter nutrient estimates. Mesoscale variation of wintertime nitrate concentrations at Biotrans are moderate (less than ±15% of the regional mean value of about 8 μmol dm⁻³). Interannual variation of the regional mean is small, too. In the available dataset, there was only 1 year with a significantly lower regional mean winter nitrate concentration (7 μmol dm⁻³), presumably due to restricted deep mixing during an atypically warm winter. The significance of winter nitrate estimates for the assessment of spring-bloom new production and the interpretation of bloom dynamics is evaluated. Applying estimates of wintertime nitrate concentrations of this study, it is found that pre-bloom new production (0.275 mol N m⁻²) at Biotrans almost equals spring-bloom new production (0.3 mol N m⁻²). Using previous estimates of wintertime nitrate yields unrealistically high estimates of pre-bloom new production (1.21–1.79 mol N m⁻²) which are inconsistent with observed levels of primary production and the seasonal development of biomass.     

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.     

Uptake of nitrogenous nutrients by phytoplankton in a barrier Island estuary: Great South Bay,New York

The uptake of urea, nitrate and ammonium by phytoplankton was measured using ¹⁵N isotopes over a one-year period in Great South Bay, a shallow coastal lagoon. The bay is a unique environment for the study of nutrient uptake since ambient concentrations of $\text{NO}{}_3{}^{\mathrm{?}}\text{NH}{}_4{}^{\mathrm{+}}$ and urea remain relatively high through the year, and phytoplankton are probably never nutrient limited. Urea nitrogen averaged 52% of the total assimilated, while ammonium represented 33% and nitrate 13%. High rates of ammonium uptake occurred only at low urea concentrations (ca< 1-μg-atom urea l^?1). Over the sampling period urea was present in relatively high concentrations, averaging 5·35 μg-atom N l^?1, while means for ammonium and nitrate averaged 1·94 and 0·65 μg-atom N l^?1, respectively. Total N uptake measured with ¹⁵N averaged about 3·3 times the calculated (from elemental ratios and ¹⁴C productivity measurements) N needs of the phytoplankton population. Highest nitrogen uptake occurred in the summer and coincided with the primary production maximum.     

Air–sea carbon dioxide fluxes in the coastal southeastern tropical Pacific

Comprehensive sea surface surveys of the partial pressure of carbon dioxide (pCO₂) have been made in the upwelling system of the coastal (0–200 km from shore) southeastern tropical Pacific since 2004. The shipboard data have been supplemented by mooring and drifter based observations. Air–sea flux estimates were made by combining satellite derived wind fields with the direct sea surface pCO₂ measurements. While there was considerable spatial heterogeneity, there was a significant flux of CO₂ from the ocean to the atmosphere during all survey periods in the region between 4° and 20° south latitude. During periods of strong upwelling the average flux out of the ocean exceeded 10 moles of CO₂ per square meter per year. During periods of weaker upwelling and high productivity the CO₂ evasion rate was near 2.5 mol/m²/yr. The average annual fluxes exceed 5 mol/m²/yr. These findings are in sharp contrast to results obtained in mid-latitude upwelling systems along the west coast of North America where the average air–sea CO₂ flux is low and can often be from the atmosphere into the ocean. In the Peruvian upwelling system there are several likely factors that contribute to sea surface pCO₂ levels that are well above those of the atmosphere in spite of elevated primary productivity: (1) the upwelling source waters contain little pre-formed nitrate and are affected by denitrification, (2) iron limitation of primary production enhanced by offshore upwelling driven by the curl of the wind stress and (3) rapid sea surface warming. The combined carbon, nutrient and oxygen dynamics of this region make it a candidate site for studies of global change.