Nutrient controls on new production in the Bodega Bay, California, coastal upwelling plume

A theoretical framework for the time-dependent processes leading to the high rates of new production in eastern boundary upwelling systems has been assembled from a series of past upwelling studies. As part of the CoOP WEST (Wind Events and Shelf Transport) study, new production in the Bodega Bay upwelling area and it's control by ambient nitrate and ammonium concentrations and the advective wind regime are described. Data and analyses are focused primarily on the WEST 2001 cruise (May–June 2001) when the two legs differed greatly in wind regimes but not nutrient concentrations. Elevated concentrations of ammonium in upwelled water with high nitrate were observed in both legs. Nitrate uptake by phytoplankton as a function of nitrate concentration was linear rather than Michaelis–Menten-like, modulated by inhibitory levels of ammonium, yielding coefficients that enable the specific nitrate uptake element of new production to be estimated from nutrient concentrations. The range of specific nitrate uptake rates for the two legs of WEST 2001 were similar, essentially a physiological response to nutrient conditions. However, the low “realization” of new production i.e. incorporation of biomass as particulate nitrogen that occurred in this system compared to the theoretical maximum possible was determined by the strong advective and turbulent conditions that dominated the second leg of the WEST 2001 study. These data are compared with other upwelling areas using a physiological shift-up model [Dugdale, R.C., Wilkerson, F.P., Morel, A. 1990. Realization of new production in coastal upwelling areas: a means to compare relative performance. Limnology and Oceanography 35, 822–829].     

Local remineralization patterns in the mesopelagic zone of the Eastern North Atlantic, off the NW Iberian Peninsula

The short-time-scale variability of the remineralization patterns in the domain of Eastern North Atlantic Central Waters (ENACW) off the NW Iberian Peninsula is studied based on biogeochemical data (oxygen, nutrient salts, total alkalinity, pH, dissolved organic matter and fluorescence of dissolved humic substances) collected weekly between May 2001 and April 2002. The temporal variability of inorganic variables points to an intensification of remineralization during the summer and autumn, with an increase of nutrients, total inorganic carbon and fluorescence and a decrease of oxygen. During the subsequent winter mixing, there is a biogeochemical reset of the system, with lower nutrients, total inorganic carbon and fluorescence and higher oxygen. In contrast to inorganic variables, the levels of dissolved organic matter in the ENACW seem to respond to short-term events probably associated with fast sinking particles, where solubilisation of organic matter prevails over remineralization. Applying a previously published stoichiometric model, we observed a vertical fractionation of organic-matter remineralization. Although there is a preferential remineralization of proteins and P compounds in the entire domain of ENACW, the percentage was higher in the upper ENACW (σ<27.10 kg/m³) than in the lower; the percentage of N and P compounds in the oxidised organic matter was >80% for the upper ENACW and 63% for the lower. Likewise, the redissolution of calcareous structures contributes about 6% and 13% to the carbon regenerated in the upper and lower layers of ENACW, respectively.     

Genera as proxies for species α‐ and β‐diversity: tested across a deep‐water Atlantic–Arctic boundary

The deep‐sea floor is increasingly subject to anthropogenic impacts. Consequently, there are increasing efforts to develop appropriate management strategies. Species‐level indicators and assessments are hampered in the deep sea by the high proportion of unknown species routinely encountered. If environmental management is to keep pace with exploitation, alternative approaches including higher taxon surrogacy (taxonomic sufficiency) must be considered. Here we compare genus‐ and species‐level studies of the diversity and ecology of deep‐sea macrobenthos on the West Shetland Slope (NE Atlantic). This is an environmentally complex region that encompasses a biogeographic transition from temperate North Atlantic to Arctic conditions, and so may be particularly challenging for any general relationship between species‐ and genus‐level analyses. We ask two questions: (i) does genus diversity reflect species diversity and (ii) does genus‐level ecology reflect species‐level ecology? We conclude that among the West Shetland Slope macrobenthos: (i) genus‐level α‐ and β‐diversity measures are highly correlated with and good predictors of their species‐level equivalents and (ii) that their ecology is very well described by genus‐level data. We further note that, given the complexity of the West Shetland Slope environment, it may be reasonable to expect these conclusions to hold in other deep‐sea environments.     

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.     

The alpha/beta ocean distinction: A perspective on freshwater fluxes, convection, nutrients and productivity in high-latitude seas

Stratification is perhaps the most important attribute of oceans with regards to climate and biology. Two simple aspects of the ocean's climate system appear to have a surprisingly important role in transforming waters that feed the global thermohaline circulation, dominating patterns of biogeochemical flux and establishing macroecological domains. First, largely because of meridional distillation (mainly due to the atmospheric transport of freshwater across the Isthmus of Panama) the North Pacific is fresher than the North Atlantic. Second, largely because of zonal distillation (e.g., warming and evaporation at low latitudes and poleward transport of latent heat and moisture by the atmosphere) the upper layers of subtropical seas are permanently stratified by temperature (N_T²=gαdT/dz>0; here called alpha oceans), while the upper layers of high-latitude seas are permanently stratified by salinity (N_S²=gβdS/dz>0; here called beta oceans). The physical basis for the boundary separating alpha and beta oceans is unclear, but may lie in the thermodynamical equations published by Fofonoff [1961. Energy transformations in the sea. Fisheries Research Board of Canada, Report Series 109, 82pp]. Nevertheless, it is clear that the resulting thermohaline distributions establish a ‘downhill journey’ of low-salinity (and nutrient-rich) waters from the North Pacific to the Arctic and then into the North Atlantic. The Arctic Ocean—itself—acts a double estuary, whereby waters entering from the North Atlantic become either denser through cooling (negative estuary) or lighter by freshening (positive estuary) as they circulate within the basin and then return to the North Atlantic as a variety of components of the ocean's conveyor. Intermediate and deep waters generally form within cyclonic beta oceans in close proximity to alphas systems. Similar patterns of stratification, nutrients and biogeographical boundaries persist in the Southern Hemisphere. It is thus argued that this simple distinction—alpha versus beta oceans—provides a broad, conceptual framework for simple interpretation of key physical and biological processes and rates, including the impacts of climate variability.     

Time-series observation of dissolved inorganic carbon and nutrients in the northwestern North Pacific

Time-series measurements of dissolved inorganic carbon (DIC) and nutrient concentrations were conducted in the northwestern North Pacific from October 2002 to August 2004. Assuming that data obtained in different years represented time-series seasonal data for a single year, vertical distributions of DIC and nutrients showed large seasonal variabilities in the surface layer (∼100 m). Seasonal variabilities in normalized DIC (nDIC) and nitrate concentrations at the sea surface were estimated to be 81–113 μmol kg⁻¹ and 12.7–15.7 μmol kg⁻¹, respectively, in the Western Subarctic Gyre. The variability in nutrients between May and July was generally at least double that in other seasons. In the Western Subarctic Gyre, estimations based on statistical analyses revealed that seasonal new production was 39–61 gC m⁻² and tended to be higher in the southwestern regions or coastal regions. The seasonal new productions in the northwestern North Pacific were two or more times higher than in the North Pacific subtropical gyre and the northeastern North Pacific. It is likely that this difference is due to spatial variations in the concentrations of trace metals and the species of phytoplankton present. In addition, from estimations of surface pCO₂ it was verified that the Western Subarctic Gyre is a source of atmospheric CO₂ between February and May and a sink for CO₂ between July and October.     

A comparative analysis of recruitment variability in North Atlantic flatfishes — testing the species range hypothesis

The hypothesis that recruitment variation in flatfishes should be most variable at the northern edge of the species range, least near the centre of the range, and intermediate near the southern limit was tested using stock and recruitment data generated from sequential population analysis for several different flatfish stocks involving four species (plaice Pleuronectes platessa, sole Solea vulgaris from the eastern Atlantic, American plaice Hippoglossoides platessoides, and yellowtail flounder Limanda ferruginae from the western Atlantic). Several groundfish species have been found to conform to this so-called species range hypothesis with the suggestion that density-independent processes predominate at the edges of the distributional range and density-dependent processes dominate in the centre of the range. Our results were generally inconsistent with the hypothesis: the coefficient of variation (CV) of recruitment for plaice in the eastern Atlantic was independent of latitude, the CV of recruitment for sole exhibited a dome-shaped relationship with latitude with the highest CVs occurring at the mid-point of the range, and the CV of recruitment for the western Atlantic stocks exhibited a monotonic decrease with latitude. We extended our latitudinal analyses by assessing both the degree of dependency of recruitment on spawning stock biomass and the spatial and temporal scales of variability in recruitment and pre-recruit survival for the eastern Atlantic stocks. In general our analysis revealed no evidence of a strong stock and recruitment relationship for any of the stocks examined, and previously published analyses revealed no such patterns with latitude. Analysis of both de-trended recruitment and pre-recruit survival time series over the species ranges of sole and plaice revealed strong positive correlations among adjacent stocks and inverse correlations among stocks at the extremes of the range. Recruitment variation in the flatfish stocks examined appears to be dominated by density-independent factors, operating at a local scale, on the egg and larval stages.     

Long-term simulation of the eutrophication of the North Sea: temporal development of nutrients, chlorophyll and primary production in comparison to observations

The ecosystem model ERSEM II has been used to hindcast the development of the ecosystem of the North Sea during the years 1955 to 1993. The simulation was driven by the box-aggregated output from a general circulation model of the North Sea of corresponding duration; radiation, river inputs, atmospheric input and boundary conditions at the borders to the Atlantic Ocean and to the Baltic Sea were applied as realistically as possible. The general features of the eutrophication process are reproduced in the hindcast: the coastal areas show strong changes in nutrient concentrations in the hindcast as well as in the observations. Eutrophication not only shows up in the nutrient concentrations, but also in primary production. The simulated spatial distributions of phosphate, nitrate and primary production compare well with the observed ones. In addition, the hindcast simulates considerable trend-like changes of the nutrients in the southern part of the North Sea, where the nutrients are transported from the continental coastal strip to the southern central North Sea. The line from the river Humber to southern Norway separates the region of noticeable anthropogenic influence of riverine and atmospheric input from the northern area, which is mainly influenced by the Atlantic nutrient inflow. The observed annual cycles in the central and northern North Sea are quite well reproduced by the hindcast. The comparison of the hindcast with the long-term observations at two sites in the continental coastal zone of the North Sea shows that the long-term behaviour of phosphate, nitrate and silicate is simulated well. Primary production is increased in summers during the main period of eutrophication, 1975 to 1989, in the hindcast and in the observations. The flagellates at Helgoland, however, experience much more pronounced annual cycles with much less interannual variability in the hindcast than in the observations.     

Benthic foraminifera and sediment grain size variability at intermediate water depths in the Northeast Atlantic during the late Pliocene–early Pleistocene

Sedimentological and faunal records from the transitional period marking the onset of widespread northern hemisphere glaciation have been investigated at Ocean Drilling Program Site 984. The late Pliocene interglacial sediments of the northeast Atlantic are carbonate rich and show evidence of vigorous bottom water circulation at intermediate water depths. Contrasting this, the late Pliocene glacial sediments are characterised by carbonate dissolution and slower bottom current velocities. Weak or “leaky” Norwegian Sea overflows, undersaturated with respect to carbonate, influenced this region during the late Pliocene glacials. The early Pleistocene pattern of intermediate water circulation appears to have changed radically in the northeast Atlantic. At this time, interglacial carbonate values and inferred bottom current velocities are low. This suggests slow-flowing, undersaturated Norwegian Sea water bathing the site. The overflow increased during the early Pleistocene interglacials as the exchange between the Atlantic and Norwegian-Greenland Seas improved. The most significant feature of the early Pleistocene glacials is the increase in inferred bottom current velocity. These changes reflect a switch in deep North Atlantic convection to shallower depths during glacial periods, possibly in a manner similar to the increasing contribution of glacial intermediate water to the North Atlantic during the late Pleistocene glacials. Our results suggest that the late Pleistocene climate variability of the North Atlantic is a pervasive feature of the late Pliocene–early Pleistocene record.     

Hydrographic changes in the Labrador Sea, 1960–2005

The Labrador Sea has exhibited significant temperature and salinity variations over the past five decades. The whole basin was extremely warm and salty between the mid-1960s and early 1970s, and fresh and cold between the late 1980s and mid-1990s. The full column salinity change observed between these periods is equivalent to mixing a 6 m thick freshwater layer into the water column of the early 1970s. The freshening and cooling trends reversed in 1994 starting a new phase of heat and salt accumulation in the Labrador Sea sustained throughout the subsequent years. It took only a decade for the whole water column to lose most of its excessive freshwater, reinstate stratification and accumulate enough salt and heat to approach its record high salt and heat contents observed between the late 1960s and the early 1970s. If the recent tendencies persist, the basin’s storages of salt and heat will fairly soon, likely by 2008, exceed their historic highs.The main process responsible for the net cooling and freshening of the Labrador Sea between 1987 and 1994 was deep winter convection, which during this period progressively developed to its record depths. It was caused by the recurrence of severe winters during these years and in its turn produced the deepest, densest and most voluminous Labrador Sea Water (LSW_1987–1994) ever observed. The estimated annual production of this water during the period of 1987–1994 is equivalent to the average volume flux of about 4.5 Sv with some individual annual rates exceeding 7.0 Sv. Once winter convection had lost its strength in the winter of 1994–1995, the deep LSW_1987–1994 layer lost “communication” with the mixed layer above, consequently losing its volume, while gaining heat and salt from the intermediate waters outside the Labrador Sea.While the 1000–2000 m layer was steadily becoming warmer and saltier between 1994 and 2005, the upper 1000 m layer experienced another episode of cooling caused by an abrupt increase in the air-sea heat fluxes in the winter of 1999–2000. This change in the atmospheric forcing resulted in fairly intense convective mixing sufficient to produce a new prominent LSW class (LSW₂₀₀₀) penetrating deeper than 1300 m. This layer was steadily sinking or deepening over the years following its production and is presently overlain by even warmer and apparently less dense water mass, implying that LSW₂₀₀₀ is likely to follow the fate of its deeper precursor, LSW_1987–1994. The increasing stratification of the intermediate layer implies intensification in the baroclinic component of the boundary currents around the mid-depth perimeter of the Labrador Sea.The near-bottom waters, originating from the Denmark Strait overflow, exhibit strong interannual variability featuring distinct short-term basin-scale events or pulses of anomalously cold and fresh water, separated by warm and salty overflow modifications. Regardless of their sign these anomalies pass through the abyss of the Labrador Sea, first appearing at the Greenland side and then, about a year later, at the Labrador side and in the central Labrador Basin.The Northeast Atlantic Deep Water (2500–3200 m), originating from the Iceland–Scotland Overflow Water, reached its historically freshest state in the 2000–2001 period and has been steadily becoming saltier since then. It is argued that LSW_1987–1994 significantly contributed to the freshening, density decrease and volume loss experienced by this water mass between the late 1960s and the mid 1990s via the increased entrainment of freshening LSW, the hydrostatic adjustment to expanding LSW, or both.     

Historical storminess and climate ‘see-saws’ in the North Atlantic region

The existence of a well-defined climate ‘see-saw’ across the North Atlantic region and surrounding areas has been known for over 200 years. The occurrence of severe winters in western Greenland frequently coincides with mild winters in northern Europe. Conversely, mild winters in western Greenland are frequently associated with cold winters across northern Europe. Whereas this ‘see-saw’ is normally discussed in terms of air temperature and pressure differences, here we explore how the climate ‘see-saw’ is reflected in records of historic storminess from Scotland, NW Ireland and Iceland. It is concluded that the stormiest winters in these regions during the last ca. 150 years have occurred when western Greenland temperatures have been significantly below average. In contrast, winters of reduced storminess have coincided with winters when air temperatures have been significantly above average in western Greenland. This reconstruction of winter storminess implies a relationship between chronologies of coastal erosion and the history of North Atlantic climate ‘see-saw’ dynamics with sustained winter storminess, and hence increased coastal erosion, taking place when the Icelandic low pressure cell is strongly anchored within the circulation of the northern hemisphere. Considered over the last ca. 2000 years, it would appear that winter storminess and climate-driven coastal erosion was at a minimum during the Medieval Warm Period. By contrast, the time interval from ca. AD 1420 until present has been associated with sustained winter storminess across the North Atlantic that has resulted in accelerated coastal erosion and sand drift.     

Primary production and implications for metabolic balance in Hawaiian lee eddies

Recent discrepancies between geochemical and biological approaches for determining whether ocean ecosystems are net heterotrophic or net autotrophic have led to uncertainty in the net metabolic state of open ocean ecosystems. Geochemical approaches indicate that the oceans are net positive autotrophic, but direct observations based on short-term incubation techniques suggest that the ocean is in a state of net heterotrophy. One hypothesis for the apparent discrepancy is that net autotrophic production occurs in aperiodic “bursts,” which are superimposed on a more constant background state of net heterotrophy. Mixing events, which introduce new nutrients to the surface ocean, provide one mechanism for fueling such aperiodic bursts of net production. In conjunction with the Eddy Flux (E-Flux) program in the lee of the Hawaiian Islands during winter 2004–2005, we examined the relationship between photosynthesis and irradiance (P vs. E) in surface waters inside and outside of two cold-core, cyclonic eddies, and conducted five incubation experiments to examine the metabolic response of mixed-layer plankton communities to nutrient-rich deep-sea water additions. Our results showed that in the mixed layer, maximum rates of light-saturated photosynthesis, derived from photosynthesis–irradiance experiments were not significantly different inside vs. outside the eddies (p=0.35 and 0.44 for E-Flux I and E-Flux III, respectively). Addition of nutrients to mixed-layer water showed that (1) gross primary production (GPP) became decoupled from a more constant rate of respiration and (2) net system metabolism shifted from approximate balance, or slight net heterotrophy, to a demonstrably net autotrophic system. From these results, we determined that the threshold GPP for net autotrophic production for the mixed layer of the study region was 1.65 mmol O₂ m⁻³ d⁻¹, which is consistent with previous estimates for the oligotrophic open ocean.     

厦门湾上屿附近海域初级生产力及相关要素的时间系列

1996年9－10月在厦门湾上屿附近近海域定点站连续6个航次测定了初级生产力和其它水文、化学和生物学要素,在较短的时间尺度内研究了初级生产力及相关要素的时间变异特征,初级生产力,叶绿素a,颗粒有机碳（POC）,总悬浮颗粒物（TSM）和营养盐等要素随时间的变化表明了沿岩水域海洋过程发生的短暂性和沿岸水体中颗粒物质来源的复杂性。     

Production and respiration control the marine microbial metabolic balance in the eastern North Atlantic subtropical gyre

Two main contrasted hypotheses have arisen during the last decades about the factors controlling the planktonic net metabolic balance in oligotrophic waters: gross primary production controls net community production vs. variability of net community production is also influenced by changes in microbial respiration. This work discusses both hypotheses analyzing the variability of metabolic rates along a gradient from the margin to the centre of the North Atlantic oligotrophic gyre, i.e. from relatively productive to more oligotrophic conditions. Net community production (NCP) was close to zero (between −3.34 and −11.77 mmol O₂ m⁻² d⁻¹) at the margin of the gyre and tended towards net heterotrophy (−44.03 mmol O₂ m⁻² d⁻¹) to the centre of the gyre as both gross primary production (GPP) and community respiration (CR) decreased. The strong relationships found between nutrient availability and both NCP and GPP suggest that factors controlling GPP are prevalent in determining NCP variability in this biogeographic region. However implementation of existing models to predict NCP from the measured GPP indicates that the precise estimation of NCP in different oligotrophic systems requires consideration of the magnitude and variability of microbial respiration rates.     

The Continuous Plankton Recorder survey and the North Atlantic Oscillation: Interannual- to Multidecadal-scale patterns of phytoplankton variability in the North Atlantic Ocean

At interannual to multidecadal time scales, much of the oceanographic and climatic variability in the North Atlantic Ocean can be associated with the North Atlantic Oscillation (NAO). While evidence suggests that there is a relationship between the NAO and zooplankton dynamics in the North Atlantic Ocean, the phytoplankton response to NAO-induced changes in the environment is less clear. Time series of monthly mean phytoplankton colour values, as compiled by the Continuous Plankton Recorder (CPR) survey, are analysed to infer relationships between the NAO and phytoplankton dynamics throughout the North Atlantic Ocean. While a few areas display highly significant (p < 0.05) trends in the CPR colour time series during the period 1948–2000, nominally significant (p < 0.20) positive trends are widespread across the basin, particularly on the continental shelves and in a transition zone stretching across the Central North Atlantic. When long-term trends are removed from both the NAO index and CPR colour time series, the correlation between them ceases to be significant. Several hypotheses are proposed to explain the observed variability in the CPR colour and its relationship with climate in the North Atlantic.     

珠江口初级生产力和新生产力研究

1996年12月和1997年8月在珠江河口湾及其毗邻海域对浮游植物生物量、初级生产力和新生产力及其环境制约机制的研究.结果表明,调查海区的叶绿素a、初级生产力和新生产力均是夏季高于冬季,冬、夏两季的平均值分别为(0.95±0.41)和(1.08±0.52)μg/dm3,(69.2±75.5)和(198.7±119.1)mg/(m2·d),(1.46±0.79)和(3.05±3.09)mg/(m3·h).冬、夏两季平均f-比分别为0.45和0.38.分级叶绿素a结果显示,冬、夏两季均以微型和微微型级分(<20μm)占优势,其对海区叶绿素a的贡献分别为796%和81.6%,对初级生产力的贡献分别为70.7%和896%.调查海区具显著的空间区域化特征,叶绿素a和潜在初级生产力的高值出现在冲淡水区的中部,向口门区和远岸区逐渐降低.现场初级生产力的高值出现在远岸区,它与复合参数B_eZ_pI₀(B_e为真光层平均叶绿素a,Z_p为真光层深度,I₀为海面光辐射强度-PAR)呈很好的正相关,说明光是研究海区初级生产力的主要限制因子.新生产力冬、夏两季的高值分别出现在交椅湾和伶仃洋西南部.     

Oil and fishing — conflict in the North Sea

A programme of research is underway at Aberdeen University on the conflicts arising in the North Sea between the fishing and offshore oil and gas industries. This research note summarizes the work that has been undertaken to date and the main aspects on which future work will concentrate.