The significance of nitrogen regeneration for new production within a filament of the Mauritanian upwelling system

The Lagrangian progression of a biological community was followed in a filament of the Mauritanian upwelling system, north-west Africa, during offshore advection. The inert dual tracers sulfur hexafluoride and helium-3 labelled a freshly upwelled patch of water that was mapped for 8 days. Changes in biological, physical, and chemical characteristics were measured, including phytoplankton productivity, nitrogen assimilation, and regeneration. Freshly upwelled water contained high nutrient concentrations but was depleted in N compared to Redfield stoichiometry. The highest rate of primary productivity was measured on the continental shelf, associated with high rates of nitrogen assimilation and a phytoplankton community dominated by diatoms and flagellates. Indicators of phytoplankton abundance and activity decreased as the labelled water mass transited the continental shelf slope into deeper water, possibly linked to the mixed layer depth exceeding the light penetration depth. By the end of the study, the primary productivity rate decreased and was associated with lower rates of nitrogen assimilation and lower nutrient concentrations. Nitrogen regeneration and assimilation took place simultaneously. Results highlighted the importance of regenerated NHC 4 in sustaining phytoplankton productivity and indicate that the upwelled NO3 pool contained an increasing fraction of regenerated NO3 as it advected offshore. By calculating this fraction and incorporating it into an f ratio formulation, we estimated that of the 12:38Tg C of annual regional production, 4:73Tg C was exportable.

The significance of nitrogen regeneration for new production within a filament of the Mauritanian upwelling system

The Lagrangian progression of a biological community was followed in a filament of the Mauritanian upwelling system, north-west Africa, during offshore advection. The inert dual tracers sulfur hexafluoride and helium-3 labelled a freshly upwelled patch of water that was mapped for 8 days. Changes in biological, physical, and chemical characteristics were measured, including phytoplankton productivity, nitrogen assimilation, and regeneration. Freshly upwelled water contained high nutrient concentrations but was depleted in N compared to Redfield stoichiometry. The highest rate of primary productivity was measured on the continental shelf, associated with high rates of nitrogen assimilation and a phytoplankton community dominated by diatoms and flagellates. Indicators of phytoplankton abundance and activity decreased as the labelled water mass transited the continental shelf slope into deeper water, possibly linked to the mixed layer depth exceeding the light penetration depth. By the end of the study, the primary productivity rate decreased and was associated with lower rates of nitrogen assimilation and lower nutrient concentrations. Nitrogen regeneration and assimilation took place simultaneously. Results highlighted the importance of regenerated NHC 4 in sustaining phytoplankton productivity and indicate that the upwelled NO3 pool contained an increasing fraction of regenerated NO3 as it advected offshore. By calculating this fraction and incorporating it into an f ratio formulation, we estimated that of the 12:38Tg C of annual regional production, 4:73Tg C was exportable.

Drivers of euphausiid species abundance and numerical abundance in the Atlantic Ocean

Mid-ocean ridges are common features of the world’s oceans but there is a lack of understanding as to how their presence affects overlying pelagic biota. The Mid-Atlantic Ridge (MAR) is a dominant feature of the Atlantic Ocean. Here, we examined data on euphausiid distribution and abundance arising from several international research programmes and from the continuous plankton recorder. We used a generalized additive model (GAM) framework to explore spatial patterns of variability in euphausiid distribution on, and at either side of, the MAR from 60°N to 55°S in conjunction with variability in a suite of biological, physical and environmental parameters. Euphausiid species abundance peaked in mid-latitudes and was significantly higher on the ridge than in adjacent waters, but the ridge did not influence numerical abundance significantly. Sea surface temperature (SST) was the most important single factor influencing both euphausiid numerical abundance and species abundance. Increases in sea surface height variance, a proxy for mixing, increased the numerical abundance of euphausiids. GAM predictions of variability in species abundance as a function of SST and depth of the mixed layer were consistent with present theories, which suggest that pelagic niche availability is related to the thermal structure of the near surface water: more deeply-mixed water contained higher euphausiid biodiversity. In addition to exposing present distributional patterns, the GAM framework enables responses to potential future and past environmental variability including temperature change to be explored.

Marine snow studies in the Northeast Atlantic Ocean: distribution, composition and role as a food source for migrating plankton

During a 25 d Lagrangian study in May and June 1990 in the Northeast Atlantic Ocean, marine snow aggregates were collected using a novel water bottle, and the composition was determined microscopically. The aggregates contained a characteristic signature of a matrix of bacteria, cyanobacteria and autotrophic picoplankton with inter alia inclusions of the tintiniid Dictyocysta elegans and large pennate diatoms. The concentration of bacteria and cyanobacteria was much greater on the aggregates than when free-living by factors of 100 to 6000 and 3000 to 2 500 000, respectively, depending on depth. Various species of crustacean plankton and micronekton were collected, and the faecal pellets produced after capture were examined. These often contained the marine snow signature, indicating that these organisms had been consuming marine snow. In some cases, marine snow material appeared to dominate the diet. This implies a food-chain short cut wherby material, normally too small to be consumed by the mesozooplankton, and considered to constitute the diet of the microplankton can become part of the diet of organisms higher in the food-chain. The micronekton was dominated by the amphipod Themisto compressa, whose pellets also contained the marine snow signature. Shipboard incubation experiments with this species indicated that (1) it does consume marine snow, and (2) its gut-passage time is sufficiently long for material it has eaten in the upper water to be defecated at its day-time depth of several hundred meters. Plankton and micronekton were collected with nets to examine their vertical distribution and diel migration and to put into context the significance of the flux of material in the guts of migrants. “Gut flux” for the T. compressa population was calculated to be up to 2% of the flux measured simultaneously by drifting sediment traps and <5% when all migrants are considered. The in situ abundance and distribution of marine snow aggregates (>0.6 mm) was examined photographically. A sharp concentration peak was usually encountered in the depth range 40 to 80 m which was not associated with peaks of in situ fluorescence or attenuation but was just below or at the base of the upper mixed layer. The feeding behaviour of zooplankton and nekton may influence these concentration gradients to a considerable extent, and hence affect the flux due to passive settling of marine snow aggregates.

Warmer, deeper and greener mixed layers in the north Atlantic subpolar gyre over the last 50 years.

Shifts in global climate resonate in plankton dynamics, biogeochemical cycles, and marine food webs. We studied these linkages in the North Atlantic subpolar gyre (NASG), which hosts extensive phytoplankton blooms. We show that phytoplankton abundance increased since the 1960s in parallel to a deepening of the mixed layer and a strengthening of winds and heat losses from the ocean, as driven by the low frequency of the North Atlantic Oscillation (NAO). In parallel to these bottom-up processes, the top-down control of phytoplankton by copepods decreased over the same time period in the western NASG, following sea surface temperature changes typical of the Atlantic Multi-decadal Oscillation (AMO). While previous studies have hypothesized that climate-driven warming would facilitate seasonal stratification of surface waters and long-term phytoplankton increase in subpolar regions, here we show that deeper mixed layers in the NASG can be warmer and host a higher phytoplankton biomass. These results emphasize that different modes of climate variability regulate bottom-up (NAO control) and top-down (AMO control) forcing on phytoplankton at decadal timescales. As a consequence, different relationships between phytoplankton, zooplankton, and their physical environment appear subject to the disparate temporal scale of the observations (seasonal, interannual, or decadal). The prediction of phytoplankton response to climate change should be built upon what is learnt from observations at the longest timescales.

Warmer, deeper and greener mixed layers in the north Atlantic subpolar gyre over the last 50 years.

Shifts in global climate resonate in plankton dynamics, biogeochemical cycles, and marine food webs. We studied these linkages in the North Atlantic subpolar gyre (NASG), which hosts extensive phytoplankton blooms. We show that phytoplankton abundance increased since the 1960s in parallel to a deepening of the mixed layer and a strengthening of winds and heat losses from the ocean, as driven by the low frequency of the North Atlantic Oscillation (NAO). In parallel to these bottom-up processes, the top-down control of phytoplankton by copepods decreased over the same time period in the western NASG, following sea surface temperature changes typical of the Atlantic Multi-decadal Oscillation (AMO). While previous studies have hypothesized that climate-driven warming would facilitate seasonal stratification of surface waters and long-term phytoplankton increase in subpolar regions, here we show that deeper mixed layers in the NASG can be warmer and host a higher phytoplankton biomass. These results emphasize that different modes of climate variability regulate bottom-up (NAO control) and top-down (AMO control) forcing on phytoplankton at decadal timescales. As a consequence, different relationships between phytoplankton, zooplankton, and their physical environment appear subject to the disparate temporal scale of the observations (seasonal, interannual, or decadal). The prediction of phytoplankton response to climate change should be built upon what is learnt from observations at the longest timescales.

Natural iron fertilization of the Atlantic sector of the Southern Ocean by continental shelf sources of the Antarctic Peninsula

In large parts of the Southern Ocean, primary production is limited due to shortage of iron (Fe). We measured vertical Fe profiles in the western Weddell Sea, Weddell-Scotia Confluence, and Antarctic Circumpolar Current (ACC), showing that Fe is derived from benthic Fe diffusion and sediment resuspension in areas characterized by high turbulence due to rugged bottom topography. Our data together with literature data reveal an exponential decrease of dissolved Fe (DFe) concentrations with increasing distance from the continental shelves of the Antarctic Peninsula and the western Weddell Sea. This decrease can be observed 3500 km eastward of the Antarctic Peninsula area, downstream the ACC. We estimated DFe summer fluxes into the upper mixed layer of the Atlantic sector of the Southern Ocean and found that horizontal advection dominates DFe supply, representing 54 ± 15% of the total flux, with significant vertical advection second most important at 29 ± 13%. Horizontal and vertical diffusion are weak with 1 ± 2% and 1 ± 1%, respectively. The atmospheric contribution is insignificant close to the Antarctic continent but increases to 15 ± 10% in the remotest waters (>1500 km offshore) of the ACC. Translating Southern Ocean carbon fixation by primary producers into biogenic Fe fixation shows a twofold excess of new DFe input close to the Antarctic continent and a one-third shortage in the open ocean. Fe recycling, with an estimated “fe” ratio of 0.59, is the likely pathway to balance new DFe supply and Fe fixation.

Methanol acetaldehyde and acetone in the surface waters of the Atlantic Ocean

Oceanic methanol, acetaldehyde, and acetone concentrations were measured during an Atlantic Meridional Transect (AMT) cruise from the UK to Chile (49°N to 39°S) in 2009. Methanol (48–361 nM) and acetone (2–24 nM) varied over the track with enrichment in the oligotrophic Northern Atlantic Gyre. Acetaldehyde showed less variability (3–9 nM) over the full extent of the transect. These oxygenated volatile organic compounds (OVOCs) were also measured subsurface, with methanol and acetaldehyde mostly showing homogeneity throughout the water column. Acetone displayed a reduction below the mixed layer. OVOC concentrations did not consistently correlate with primary production or chlorophyll-a levels in the surface Atlantic Ocean. However, we did find a novel and significant negative relationship between acetone concentration and bacterial leucine incorporation, suggesting that acetone might be removed by marine bacteria as a source of carbon. Microbial turnover of both acetone and acetaldehyde was confirmed. Modeled atmospheric data are used to estimate the likely air-side OVOC concentrations. The direction and magnitude of air-sea fluxes vary for all three OVOCs depending on location. We present evidence that the ocean may exhibit regions of acetaldehyde under-saturation. Extrapolation suggests that the Atlantic Ocean represents an overall source of these OVOCs to the atmosphere at 3, 3, and 1 Tg yr−1 for methanol, acetaldehyde, and acetone, respectively.

Influence of the Pacific Decadal Oscillation on phytoplankton phenology and community structure in the western North Pacific

Phytoplankton phenology and community structure in the western North Pacific were investigated for 2001–2009, based on satellite ocean colour data and the Continuous Plankton Recorder survey. We estimated the timing of the spring bloom based on the cumulative sum satellite chlorophyll adata, and found that the Pacific Decadal Oscillation (PDO)-related interannual SST anomaly in spring significantly affected phytoplankton phenology. The bloom occurred either later or earlier in years of positive or negative PDO (indicating cold and warm conditions, respectively). Phytoplankton composition in the early summer varied depending on the magnitude of seasonal SST increases, rather than the SST value itself. Interannual variations in diatom abundance and the relative abundance of non-diatoms were positively correlated with SST increases for March–April and May–July, respectively, suggesting that mixed layer environmental factors, such as light availability and nutrient stoichiometry, determine shifts in phytoplankton community structure. Our study emphasised the importance of the interannual variation in climate-induced warm–cool cycles as one of the key mechanisms linking climatic forcing and lower trophic level ecosystems.

Modelling a light-driven phytoplankton succession

We used a numerical model to investigate if and to what extent cellular photoprotective capacity accounts for succession and vertical distribution of marine phytoplankton species/groups. A model describing xanthophyll photoprotective activity in phytoplankton has been implemented in the European Regional Sea Ecosystem Model and applied at the station L4 in the Western English Channel. Primary producers were subdivided into three phytoplankton functional types defined in terms of their capacity to acclimate to different light-specific environments: low light (LL-type), high light (HL-type) and variable light (VL-type) adapted species. The LL-type is assumed to have low cellular level of xanthophyll-cycling pigments (PX) relative to the modelled photosynthetically active pigments (chlorophyll and fucoxanthin (FUCO) = PSP). The HL-type has high PX content relative to PSP while VL-type presents an intermediate PX to PSP ratio. Furthermore, the VL-type is capable of reversibly converting FUCO to PX and synthesizing new PX under high-light stress. In order to reproduce phytoplankton community succession with each of the three groups being dominant in different periods of the year, we had also to assume reduced grazing pressure on HL-adapted species. Model simulations realistically reproduce the observed seasonal patterns of pigments and nutrients highlighting the reasonability of the underpinning assumptions. Our model suggests that pigment-mediated photophysiology plays a primary role in determining the evolution of marine phytoplankton communities in the winter-spring period corresponding to the shoaling of the mixed layer and the increase of light intensity. Grazing selectivity however contributes to the phytoplankton community composition in summer.

Lagrangian evolution of DMS during the Southern Ocean gas exchange experiment: The effects of vertical mixing and biological community shift

Concentrations of dimethylsulfide (DMS) and its precursor dimethylsulfoniopropionate (DMSP) are highly variable in time and space. What is driving the variability in DMS(P), and can those variability be explained by physical processes and changes in the biological community? During the Southern Ocean Gas Exchange Experiment (SO GasEx) in the austral fall of 2008, two 3He/SF6 labeled patches were created in the surface water. SF6 and DMS were surveyed continuously in a Lagrangian framework, while direct measurements of air-sea exchange further constrained the gas budgets. Turbulent diffusivity at the base of the mixed layer was estimated from SF6 profiles and used to calculate the vertical fluxes of DMS and nutrients. Increasing mixed layer nutrient concentrations due to mixing were associated with a shift in the phytoplankton community structure, which in turned likely affected the sulfur dynamics on timescales of days. DMS concentration as well as air-sea DMS flux appeared to be decoupled from the DMSP concentration, possibly due to grazing and bacterial DMS production. Contrary to expectations, in an environment with high winds and modest productivity, physical processes (air-sea exchange, photochemistry, vertical mixing) only accounted for a small fraction of DMS loss from the surface water. Among the DMS sinks, inferred biological consumption most likely dominated during SO GasEx.

Variations in GDGT distributions through the water column in the South East Atlantic Ocean

The TetraEther indeX of 86 carbon atoms (TEX86) temperature proxy is widely used in reconstructions of past sea surface temperature. Most current calibrations are based on surface sediment distributions of the glycerol dialkyl glycerol tetraether lipids (GDGTs) that comprise TEX86 and assume that these GDGTs are exported from the upper mixed layer. However, GDGT export from deeper waters could impact sedimentary GDGT distributions and therefore TEX86 paleothermometry. Here we examine GDGT distributions in suspended particulate matter (SPM) and underlying sediments collected from the Southeast Atlantic Ocean. Our results reveal different GDGT distributions - specifically the ratio between GDGTs bearing 2 vs. 3 cyclopentyl moieties, [2/3] ratios - between surface, subsurface (>50-200 m) and deep water (>200 m) SPM, which suggests the occurrence of in situ (deep) production that is not apparent when considering TEX86. The GDGT distributions in sediments match those of subsurface waters rather than surface waters, suggesting that they have not been preferentially derived from the upper mixed layer; this is consistent with GDGT abundances being highest in shallow subsurface SPM (˜100 to 200 m). It remains unclear what governs the different [2/3] ratios throughout the water column, but it is likely related to a combination of temperature and thaumarchaeotal community structure.

Space-based lidar measurements of global ocean carbon stocks

Global ocean phytoplankton biomass (C-phyto) and total particulate organic carbon (POC) stocks have largely been characterized from space using passive ocean color measurements. A space-based light detection and ranging (lidar) system can provide valuable complementary observations for C-phyto and POC assessments, with benefits including day-night sampling, observations through absorbing aerosols and thin cloud layers, and capabilities for vertical profiling through the water column. Here we use measurements from the Cloud-Aerosol Lidar with Orthogonal Polarization (CALIOP) to quantify global C-phyto and POC from retrievals of subsurface particulate backscatter coefficients (b(bp)). CALIOP b(bp) data compare favorably with airborne, ship-based, and passive ocean data and yield global average mixed-layer standing stocks of 0.44 Pg C for C-phyto and 1.9 Pg for POC. CALIOP-based C-phyto and POC data exhibit global distributions and seasonal variations consistent with ocean plankton ecology. Our findings support the use of spaceborne lidar measurements for advancing understanding of global plankton systems.

The origin of sub-surface source waters define the sea-air flux of methane in the Mauritanian Upwelling, NW Africa

Concentrations and flux densities of methane were determined during a lagrangian study of an advective filament in the permanent upwelling region off western Mauritania. Newly upwelled waters were dominated by the presence of North Atlantic Central Water and surface CH4 concentrations of 2.2 ± 0.3 nmol L-1 were largely in equilibrium with atmospheric values, with surface saturations of 101.7 ± 14%. As the upwelling filament aged and was advected offshore, CH4 enriched South Atlantic Central Water from intermediate depths of 100 to 350m was entrained into the surface mixed layer of the filament following intense mixing associated with the shelf break. Surface saturations increased to 198.9 ± 15% and flux densities increased from a mean value over the shelf of 2.0 ± 1.1 µmol m-2d-1 to a maximum of 22.6 µmol m-2d-1. Annual CH4 emissions for this persistent filament were estimated at 0.77 ± 0.64 Gg which equates to a maximum of 0.35% of the global oceanic budget. This raises the known outgassing intensity of this area and highlights the importance of advecting filaments from upwelling waters as efficient vehicles for air-sea exchange.

High-resolution view of the spring bloom initiation and net community production in the Subantarctic Southern Ocean using glider data

In the Southern Ocean, there is increasing evidence that seasonal to subseasonal temporal scales, and meso- to submesoscales play an important role in understanding the sensitivity of ocean primary productivity to climate change. This drives the need for a high-resolution approach to re- solving biogeochemical processes. In this study, 5.5 months of continuous, high-resolution (3 h, 2 km horizontal resolution) glider data from spring to summer in the Atlantic Subantarctic Zone is used to investigate: (i) the mechanisms that drive bloom initiation and high growth rates in the region and (ii) the seasonal evolution of water column production and respiration. Bloom initiation dates were analysed in the context of upper ocean boundary layer physics highlighting sensitivities of different bloom detection methods to different environmental processes. Model results show that in early spring (September to mid-November) increased rates of net community production (NCP) are strongly affected by meso- to submesoscale features. In late spring/early summer (late-November to mid-December) seasonal shoaling of the mixed layer drives a more spatially homogenous bloom with maximum rates of NCP and chlorophyll biomass. A comparison of biomass accumulation rates with a study in the North Atlantic highlights the sensitivity of phytoplankton growth to fine-scale dynamics and emphasizes the need to sample the ocean at high resolution to accurately resolve phytoplankton phenology and improve our ability to estimate the sensitivity of the biological carbon pump to climate change.