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.     

Large bio-geographical shifts in the north-eastern Atlantic Ocean: From the subpolar gyre, via plankton, to blue whiting and pilot whales

Pronounced changes in fauna, extending from the English Channel in the south to the Barents Sea in the north-east and off Greenland in the north-west, have occurred in the late 1920s, the late 1960s and again in the late 1990s. We attribute these events to exchanges of subarctic and subtropical water masses in the north-eastern North Atlantic Ocean, associated with changes in the strength and extent of the subpolar gyre. These exchanges lead to variations in the influence exerted by the subarctic or Lusitanian biomes on the intermediate faunistic zone in the north-eastern Atlantic. This strong and persistent bottom-up bio-physical link is demonstrated using a numerical ocean general circulation model and data on four trophically connected levels in the food chain – phytoplankton, zooplankton, blue whiting, and pilot whales. The plankton data give a unique basin-scale depiction of these changes, and a long pilot whale record from the Faroe Islands offers an exceptional temporal perspective over three centuries. Recent advances in simulating the dynamics of the subpolar gyre suggests a potential for predicting the distribution of the main faunistic zones in the north-eastern Atlantic a few years into the future, which might facilitate a more rational management of the commercially important fisheries in this region.     

Variability and propagation of Labrador Sea Water in the southern subpolar North Atlantic

The variability of two modes of Labrador Sea Water (LSW) (upper and deep Labrador Sea Water) and their respective spreading in the interior North Atlantic Ocean are investigated by means of repeated ship surveys carried out along the zonal WOCE line A2/AR19 located at 43–48°N (1993–2007) and along the GOOS line at about 48–51°N (1997–2002). Hydrographic section data are complemented by temperature, salinity, and velocity time series recorded by two moorings. They have been deployed at the western flank of the Mid-Atlantic Ridge (MAR) in the Newfoundland Basin during 1996–2004. The analysis of hydrographic anomalies at various longitudes points to a gradual eastward propagation of LSW-related signals, which happens on time scales of 3–6 years from the formation region towards the MAR. Interactions of the North Atlantic Current (NAC) with the Deep Western Boundary Current (DWBC) close to Flemish Cap point to the NAC being the main distributor of the different types of LSW into the interior of the Newfoundland Basin. Comparisons between the ship data and the mooring records revealed that the mooring sites are located in a region affected by highly variable flow. The mooring time series demonstrate an elevated level of variability with eddy activity and variability associated with the NAC considerably influencing the LSW signals in this region. Hydrographic data taken from Argo profiles from the vicinity of the mooring sites turned out to mimic quite well the temporal evolution captured by the moorings. There is some indication of occasional southward flow in the LSW layer near the MAR. If this can be considered as a hint to an interior LSW-route, it is at least of minor importance in comparison to the DWBC. It acts as an important supplier for the interior North Atlantic, distributing older and recently formed LSW modes southward along the MAR.     

On the mechanisms behind salinity anomaly signals of the northern North Atlantic

Hydrographic time series from the northern North Atlantic throughout the 20th century show oscillations in temperature and salinity at more or less regular intervals. The Great Salinity Anomalies described during the 1970s [Dickson, R.R., Meincke, J., Malmberg, S.-A., Lee, A.J., 1988. The “Great Salinity Anomaly” in the North Atlantic, 1968-1982. Progress in Oceanography 20, 103-151.], during the 1980s [Belkin, I.M., Levitus, S., Antonov, J., Malmberg, S.-A., 1998. “Great Salinity Anomalies” in the North Atlantic. Progress in Oceanography 41, 1-68.], and during the 1990s [Belkin, I.M., 2004. Propagation of the “Great Salinity Anomaly” of the 1990s around the northern North Atlantic. Geophysical Research Letters 31(8), L08306, doi:10.1029/2003GL019334.] have distinct amplitudes, and all three of them were interpreted as low salinity anomalies propagating downstream through the anti-clockwise circulation system of the northern North Atlantic Ocean. Further inspection of time series from the Northeast Atlantic and the Northwest Atlantic over the past century shows, however, several other distinct negative anomalies of lesser amplitudes. Additionally, a number of high salinity anomalies can be identified. The present paper analyses further the propagation of the negative and positive anomalies and links them together. It is shown that they have varying speeds of propagation, and that the varying speeds are correlated across the North Atlantic. We propose that varying volume fluxes in and out of the Arctic Basin is the causal mechanism behind the anomaly signals, and that the North Atlantic Oscillation (NAO) partly has influence on the flux variations described. Periods of large decadal-scale amplitudes of the NAO coincide with periods of large decadal-scale oscillation in the marine climate.     

New Determinations of the Ages of Tide in the North Atlantic Ocean

Ages of tide provide relevant information about the spatial distribution of existing anomalies in the normal modes of the oceans, because a delay may be associated with bottom friction energy dissipation, closely located resonances, bathymetric gradients, or radiational effects. The determination of other parameters, such as the age of diurnal tide or age of parallax, also provide further knowledge about the ocean's hydrodynamical response to acting forces. Following the development of new ocean models and the availability of a greater amount of data, these parameters can be redetermined. We present the spatial distribution of these parameters in the Northeast Atlantic Ocean and the Mediterranean Sea, obtained from 507 stations. The results are discussed in terms of bathymetric models, coastal features, sea surface temperature, wind and other environmental factors.     

New Determinations of the Ages of Tide in the North Atlantic Ocean

Ages of tide provide relevant information about the spatial distribution of existing anomalies in the normal modes of the oceans, because a delay may be associated with bottom friction energy dissipation, closely located resonances, bathymetric gradients, or radiational effects. The determination of other parameters, such as the age of diurnal tide or age of parallax, also provide further knowledge about the ocean's hydrodynamical response to acting forces. Following the development of new ocean models and the availability of a greater amount of data, these parameters can be redetermined. We present the spatial distribution of these parameters in the Northeast Atlantic Ocean and the Mediterranean Sea, obtained from 507 stations. The results are discussed in terms of bathymetric models, coastal features, sea surface temperature, wind and other environmental factors.     

Temporal Variations in the Dissolved Nutrient Stocks in the Surface Water of the Western North Atlantic Ocean

Changes in patterns of undetectability and molar ratios of dissolved nutrients in the euphotic zone of the oligotrophic western North Atlantic Ocean were investigated utilizing the Bermuda Atlantic Time-series Study (BATS) data set of the US Joint Global Ocean Flux Study (JGOFS). Our aim was to examine the temporal dynamics of nutrient stocks over a decade (1989∼1998) and to gain insight into the interactions between the different biotic and abiotic factors underlying BATS. Patterns of nutrient undetectability clearly revealed the depleted nature of the nutrients in surface water at the BATS location, particularly phosphorous. The N:P ratio was consistently far above the nominal Redfield ratio (mean, 38.5) but was significantly lower during the 1993∼1994 period (22.1). Over the same period the proportion of samples depleted in N only increased while the proportion of samples depleted in P only decreased. This indicates an overall reduction of N relative to P in the surface water at BATS during the 1993∼1994 period, the reasons for this anomaly, though, are not clear. The correlation analysis between the biotic and abiotic variables at BATS has indicated some interesting relationships that can help understand some of the parameters affecting nutrient stocks in the euphotic zone and their consequent impacts on marine biota. Although nutrient stocks in the oligotrophic environment are limited, they might be subject to interannual variation that may become anomalous in some cases. These variations might underlay significant feedback mechanisms by affecting marine productivity, the prime factor controlling the sequestration of atmospheric CO₂ by the oceans. This revised version was published online in July 2006 with corrections to the Cover Date.     

Oxygen minimum zones in the eastern tropical Atlantic and Pacific oceans

Within the eastern tropical oceans of the Atlantic and Pacific basin vast oxygen minimum zones (OMZ) exist in the depth range between 100 and 900 m. Minimum oxygen values are reached at 300–500 m depth which in the eastern Pacific become suboxic (dissolved oxygen content <4.5 μmol kg⁻¹) with dissolved oxygen concentration of less than 1 μmol kg⁻¹. The OMZ of the eastern Atlantic is not suboxic and has relatively high oxygen minimum values of about 17 μmol kg⁻¹ in the South Atlantic and more than 40 μmol kg⁻¹ in the North Atlantic. About 20 (40%) of the North Pacific volume is occupied by an OMZ when using 45 μmol kg⁻¹ (or 90 μmol kg⁻¹, respectively) as an upper bound for OMZ oxygen concentration for ocean densities lighter than σ_θ < 27.2 kg m⁻³. The relative volumes reduce to less than half for the South Pacific (7% and 13%, respectively). The abundance of OMZs are considerably smaller (1% and 7%) for the South Atlantic and only 0% and 5% for the North Atlantic. Thermal domes characterized by upward displacements of isotherms located in the northeastern Pacific and Atlantic and in the southeastern Atlantic are co-located with the centres of the OMZs. They seem not to be directly involved in the generation of the OMZs.OMZs are a consequence of a combination of weak ocean ventilation, which supplies oxygen, and respiration, which consumes oxygen. Oxygen consumption can be approximated by the apparent oxygen utilization (AOU). However, AOU scaled with an appropriate consumption rate (aOUR) gives a time, the oxygen age. Here we derive oxygen ages using climatological AOU data and an empirical estimate of aOUR. Averaging oxygen ages for main thermocline isopycnals of the Atlantic and Pacific Ocean exhibit an exponential increase with density without an obvious signature of the OMZs. Oxygen supply originates from a surface outcrop area and can also be approximated by the turn-over time, the ratio of ocean volume to ventilating flux. The turn-over time corresponds well to the average oxygen ages for the well ventilated waters. However, in the density ranges of the suboxic OMZs the turn-over time substantially increases. This indicates that reduced ventilation in the outcrop is directly related to the existence of suboxic OMZs, but they are not obviously related to enhanced consumption indicated by the oxygen ages. The turn-over time suggests that the lower thermocline of the North Atlantic would be suboxic but at present this is compensated by the import of water from the well ventilated South Atlantic. The turn-over time approach itself is independent of details of ocean transport pathways. Instead the geographical location of the OMZ is to first order determined by: (i) the patterns of upwelling, either through Ekman or equatorial divergence, (ii) the regions of general sluggish horizontal transport at the eastern boundaries, and (iii) to a lesser extent to regions with high productivity as indicated through ocean colour data.     

Assemblages of phytoplankton pigments along a shipping line through the North Atlantic and tropical Pacific

A set of phytoplankton pigment measurements collected on eight quarterly transects from France to New Caledonia is analyzed in order to identify the main assemblages of phytoplankton and to relate their occurrence to oceanic conditions. Pigment concentrations are first divided by the sum [monovinyl chlorophyll a plus divinyl chlorophyll a] to remove the effect of biomass, and second are normalized to give an equal weight to all pigments. The resulting 17 pigments × 799 observations matrix is then classified into 10 clusters using neural methodology. Eight out of these 10 clusters have a well marked regional or seasonal character, thus evidencing adapted responses of the phytoplankton communities. The main gradient opposes two clusters with high fucoxanthin and chlorophyll c₁₊₂ in the North Atlantic in January, April and July, to three clusters in the South Pacific Subtropical Gyre with high divinyl chlorophyll a, zeaxanthin and phycoerythrin. One of the clusters in the South Pacific Subtropical Gyre has relatively high zeaxanthin and phycoerythrin contents and dominates in November and February (austral summer), while another with relatively high divinyl chlorophylls a and b dominates in May and August (austral winter). The third one in the South Pacific is characterized by high carotene concentration and its occurrence peaks in February and May. In the equatorial current system, one cluster, rich in chlorophylls b and c₁₊₂, is strictly located in a narrow zone centred at the equator, while another with relatively high violaxanthin concentration is restricted to the high nutrient - low chlorophyll waters in only the southern part of the South Equatorial Current. One cluster with relatively high prasinoxanthin content has a spatial distribution spanning the entire South Equatorial Current. Two clusters have a ubiquitous distribution: one in the equatorial Pacific, the Carribbean Sea and the North Atlantic during summer has pigment concentrations close to the average of the entire dataset, and the other in the South Pacific Subtropical Gyre, the Carribbean Sea and the North Atlantic during autumn clearly has an oligotrophic character. Many of the differences between clusters are caused by diagnostic pigments of nano- or picoflagellates. While the space and time characteristics of the clusters are well marked and might correspond to differences in physical and chemical forcing, knowledge of the ecological requirements of these flagellates is generally lacking to explain how the variability of the environment triggers these clusters.     

Constructing an idealized model of the North Atlantic Ocean using slippery sacks

This paper documents the continued development and testing of a new Lagrangian oceanic general circulation model. The slippery sacks ocean model (SSOM), which represents a body of water as a pile of conforming parcels, is improved and is used to simulate circulations in homogeneous oceans and in an idealized model of the North Atlantic Ocean.A method for including horizontal mixing in the SSOM is presented. A given sack’s nearest neighbors are identified in the positive and negative x- and y-directions, and the sack exchanges momentum and/or tracers with these neighbors. This formulation of mixing is straightforward to implement, computationally efficient, and it produces results similar to a standard Eulerian finite-difference representation of diffusion.The model’s ability to reproduce the Stommel and Munk solutions to the classical western boundary current problem is tested. When steps are taken to reduce the potential energy barrier to sacks crossing one another, the model generates circulations that are consistent with linear theory. In moderately non-linear regimes the model produces appropriate departures from linear solutions including a boundary current that continues along the northern boundary for a time.Taking advantage of the new mixing scheme and lessons learned from simulations of homogeneous oceans, the authors construct an idealized model of the North Atlantic Ocean. They compare simulations conducted with the SSOM to similar simulations conducted with the Massachusetts Institute of Technology general circulation model (MITgcm). The SSOM and the MITgcm produce similar wind-forced gyres, thermocline structure, and meridional overturning. The SSOM is also used to explore how circulations change in the limit when tracer diffusion goes to zero.     

Assessment of excess nitrate development in the subtropical North Atlantic

Geochemical estimates of N₂ fixation in the North Atlantic often serve as a foundation for estimating global marine diazotrophy. Yet despite being well-studied, estimations of nitrogen fixation rates in this basin vary widely. Here we investigate the variability in published estimates of excess nitrogen accumulation rates in the main thermocline of the subtropical North Atlantic, testing the assumptions and choices made in the analyses. Employing one of these previously described methods, modified here with improved estimates of excess N spatial gradients and ventilation rates of the main thermocline, we determine a total excess N accumulation rate of 7.8 ± 1.7 × 10¹¹ mol N yr^− 1. Contributions to excess N development include atmospheric deposition of high N:P nutrients (adding excess N at a rate of 3.0 ± 0.9 × 10¹¹ mol N yr^− 1 for  38% of the total), high N:P dissolved organic matter advected into and mineralized in the main thermocline (adding excess N at 2.2 ± 1.1 × 10¹¹ mol N yr^− 1 for  28% of the total), and, calculated by mass balance of the excess N field, N₂ fixation (adding excess N at 2.6 ± 2.2 × 10¹¹ mol N yr^− 1 for  33% of the total). Assuming an N:P of 40 and this rate of excess N accumulation due to the process, N₂ fixation in the North Atlantic subtropical gyre is estimated at  4 × 10¹¹ mol N yr^− 1. This relatively low rate of N₂ fixation suggests that i) the rate of N₂ fixation in the North Atlantic is greatly overestimated in some previous analyses, ii) the main thermocline is not the primary repository of N fixed by diazotrophs, and/or iii) the N:P ratio of exported diazotrophic organic matter is much lower than generally assumed. It is this last possibility, and our uncertainty in the N:P ratios of exported material supporting excess N development, that greatly lessens our confidence in geochemical measures of N₂ fixation.     

Nutrient distributions in baroclinic eddies of the oligotrophic North Atlantic and inferred impacts on biology

High-sensitivity (nanomolar) techniques for nitrate and phosphate were applied to study nutrient patterns in the euphotic zone of mesoscale eddies in the Sargasso Sea during the EDDIES project. Surface concentrations of nitrate plus nitrite (DNN) and phosphate (DIP) were found in the range of 1–20 nM with substantial spatial variability in the eddies, with resulting mean N:P molar ratios of 2.1. Chlorophyll biomass was well correlated with DNN but not DIP in the upper euphotic zone, suggesting N-limitation of marine phytoplankton at this time of year. Within the upper 140 m, the water column experienced a transition from a P-enriched (relative to Redfield ratio) shallow layer to a N-enriched deep layer, which may suggest downward transport and subsequent remineralization of high N/P biogenic products presumably originating from N₂ fixation. Chlorophyll biomass in the deep chlorophyll maximum of eddies was found to be tightly related to eddy–induced variability in major nutrients (N, P, Si) and nutrient stoichiometry, suggesting that the impact of eddies on biology is through control of nutrient availability. Because the eddies were likely to be in various phases of development (different degrees of both biological and physical maturity), full interpretation of eddy data and dynamics will require better coverage of a full eddy life cycle.     

北太平洋西部和印度洋东部及其邻近海域表层水体中137Cs的分布

对北太平洋西部海域、苏禄海及印尼海、中国南海、印度洋东部海域、孟加拉湾及安达曼海等表层水体中放射性核素137Cs的活度进行了测定。结果表明,上述海域表层水体中137Cs活度显示了较大的变化范围,最低值出现在南极附近的南大洋(1.1Bqm-3),较高的活度值则出现在北太平洋西部海域及中国南海(3Bqm-3)。在所研究水域范围内,137Cs活度的纬度分布特征并没有完全有效地反映出137Cs的全球理论大气沉降趋势及其纬度效应。综合本研究及Miyake等人(1988)的测定结果,我们计算出137Cs自表层海水中的析出速率在苏禄海及印尼海约为0.016/a,在孟加拉湾及安达曼海约为0.033/a,在中国南海约为0.029/a,这一结果明显低于西北太平洋日本沿海表层水体中137Cs的析出速率。这可能是因为在这些海域,横向及纵向的水体混合过程相对都较慢,而且颗粒物对137Cs的吸附析出过程也比较弱所致。     

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.     

Verification of the wind-driven transport in the North Pacific subtropical gyre using gridded wind-stress products

Wind-stress products supplied by satellite scatterometers carried the European Remote-sensing Satellite (ERS) and QuikSCAT (QSCAT), together with numerical weather predictions from the European Centre for Medium Range Weather Forecasting (ECMWF) and the National Centre for Environmental Prediction (NCEP) were used to estimate wind-driven transports of the North Pacific subtropical gyre. At 30°N, we compared the wind-driven transports with geostrophic transports calculated from World Ocean Database 2005. The wind-driven transports for QSCAT and NCEP are in good agreement with the geostrophic transport within reasonable error, except for a regional difference in the eastern part of the section. The difference in the eastern part suggests an anti-cyclonic deviation of the geostrophic transport, resulting from an anti-cyclonic anomalous flow in the surface layer. It is suggested that this anomalous flow is the Eastern Gyral, produced by the thermohaline process associated with the formation of the Eastern Subtropical Mode Water. To investigate the validity of QSCAT and NCEP data, we examined whether or not the Sverdrup transports for these products are consistent with the transport of the western boundary current estimated by past studies. The net southward transport, given by the sum of the Sverdrup transport for QSCAT and NCEP and the thermohaline transport, agrees well with the net northward transport of the western boundary current. From this result, together with the fact that the wind-driven transports for these products are in good agreement with the geostrophic transport, we conclude that the Sverdrup balance can hold in the North Pacific subtropical gyre.     

Magnetic anomalies in the canary basin and the Mesozoic evolution of the central North Atlantic

The data from a recent magnetic compilation by Verhoefet al. (1991) off west Africa were used in combination with data in the western Atlantic to review the Mesozoic plate kinematic evolution of the central North Atlantic. The magnetic profile data were analyzed to identify the M-series sea floor spreading anomalies on the African plate. Oceanic fracture zones were identified from magnetic anomalies and seismic and gravity measurements. The identified sea floor spreading anomalies on the African plate were combined with those on the North American plate to calculate reconstruction poles for this part of the central Atlantic. The total separation poles derived in this paper describe a smooth curve, suggesting that the motion of the pole through time was continuous. Although the new sea floor spreading history differs only slightly from the one presented by Klitgord and Schouten (1986), it predicts smoother flowlines. On the other hand, the sea floor spreading history as depicted by the flowlines for the eastern central Atlantic deviates substantially from that of Sundvik and Larson (1988). A revised spreading history is also presented for the Cretaceous Magnetic Quiet Zone, where large changes in spreading direction occurred, that can not be resolved when fitting magnetic isochrons only, but which are evident from fracture zone traces and directions of sea floor spreading topography.Deceased 11 November 1991