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.     

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.     

Tracing dissolved organic matter cycling in the eastern boundary of the temperate North Atlantic using absorption and fluorescence spectroscopy

Apparent oxygen utilization (AOU), dissolved organic carbon (DOC), coloured dissolved organic matter (CDOM) absorption spectra, and CDOM fluorescence characteristic of aromatic amino acids (excitation/emission 280 nm/320 nm; F(280/320)) and marine-humic like substances (320 nm/410 nm; F(320/410)) were measured in full depth during a cruise in the temperate Eastern North Atlantic (ENA). An optimum multi-parameter (OMP) inverse method was run to calculate water mass proportion-weighted average (archetypal) concentrations of these chemical parameters for all water masses and samples. Archetypal concentrations retain the variability due to water mass mixing and basin scale mineralization from the water mass formation sites to the study area. Conversely, the difference between measured and archetypal concentrations, retain the variability due to dissimilarities in mineralization processes within the study area. Our analysis indicates that DOC supported 26±3% of the AOU in the dark temperate ENA and that basin scale processes occurring at and from the formation area of the water masses explained 63% of the total DOC variability. Our data also suggests that DOC remineralized at the basin scale was of lower molecular weight, and with a lower proportion of fluorescent aromatic amino acids than found within the study area. The relationship between the absorption coefficient at 254 nm (a_CDOM(254)) and AOU indicates that a_CDOM(254) was consumed during organic matter remineralization in the dark ocean, with 55% of the variability being explained by basin scale processes. The relationships of F(320/410) with AOU and DOC confirmed that marine humic-like substances are produced by microbial degradation processes, at a rate of 6.1±0.9×10⁻³ mg equivalents of QS mol AOU⁻¹. Our results also indicate that basin-scale remineralization processes account for 85% of the total variability of F(320/410), emphasizing that large scale processes control the formation of humic-like substance in the dark ENA.     

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.     

基于WOA13数据的北大西洋声传播分析

应用WOA13季节平均数据和BELLHOP模型,在季节、声源频率、声源深度和掠射角等因素确定的情况下,分析北大西洋冬季(1-3月)声道轴深度、最小声速值、表层声速值的分布,通过仿真计算研究选用位置点5 m深度声源的声传播规律:反转深度随纬度升高而降低,低纬度海岭东西两侧差别不大,15°N以北为西侧大于东侧。55°N以南海区可形成汇聚区波导,海岭西侧的汇聚区跨度大于海岭东侧,有混合层时还存在一定强度的表面波导,汇聚区处5 m、100 m和250 m接收深度上的传播损失差异较小,增益为7～19 dB,55°N以北海区则为有焦散结构的表面波导。以北大西洋35°N为界,以南以汇聚区波导探测有利,以北以表面波导探测有利。     

Trans-Atlantic responses of Calanus finmarchicus populations to basin-scale forcing associated with the North Atlantic Oscillation

Populations of the copepod species Calanus finmarchicus often dominate the springtime biomass and secondary production of shelf ecosystems throughout the North Atlantic Ocean. Recently, it has been hypothesised that interannual to interdecadal fluctuations observed in such populations are driven primarily by climate-associated changes in ocean circulation. Here, we compare evidence from the North Sea and Gulf of Maine/Western Scotian Shelf (GoM/WSS) linking fluctuations in C. finmarchicus abundance to changes in ocean circulation associated with the North Atlantic Oscillation (NAO). A particularly striking contrast emerges from this Trans-Atlantic comparison: whereas the North Sea C. finmarchicus population exhibits a negative correlation with the NAO index, the GoM/WSS population exhibits a more complex, positive association with the index. The physical processes underlying these contrasting population responses are discussed in the context of regional- to basin-scale circulation changes associated with the NAO.     

2016与1998年春季北大西洋海表温度异常的差异及成因

利用再分析资料以及混合层海温诊断方程, 研究1997—1998与2015—2016年超级厄尔尼诺次年北大西洋海表温度异常(sea surface temperature anomalies, SSTA)的差异及成因。结果显示, 北大西洋SSTA在1998年春季呈明显正负正三极型式分布, 而在2016年呈弱的负正负型态。诊断热带北大西洋SSTA的影响因素表明, 1998年春季暖SSTA除了之前研究强调的海洋表面向大气的潜热输送异常减少, 以及吸收太阳辐射的增加外, 海洋动力过程即Ekman纬向漂流也起着重要的作用。热力过程与厄尔尼诺峰值后出现的北大西洋涛动(North Atlantic Oscillation, NAO)负位相有关, 其可引起亚速尔高压减弱, 产生西南风异常, 通过风-蒸发-海表温度(sea surface temperature, SST)反馈机制使热带北大西洋蒸发减弱, 海表增暖, 沃克环流下沉支的东移对这一增暖也有贡献。与1997—1998厄尔尼诺事件不同, 2015—2016厄尔尼诺事件没有强迫出负位相NAO, 而是出现弱NAO正位相, 热带北大西洋为弱的东风异常, 使海表发生一定的冷却, 形成2016春季北大西洋SSTA与1998年的明显差异。     

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.