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The Green Edge project was designed to investigate the onset, life, and fate of a phytoplankton spring bloom (PSB) in the Arctic Ocean. The lengthening of the ice-free period and the warming of seawater, amongst other factors, have induced major changes in Arctic Ocean biology over the last decades. Because the PSB is at the base of the Arctic Ocean food chain, it is crucial to understand how changes in the Arctic environment will affect it. Green Edge was a large multidisciplinary, collaborative project bringing researchers and technicians from 28 different institutions in seven countries together, aiming at understanding these changes and their impacts on the future. The fieldwork for the Green Edge project took place over two years (2015 and 2016) and was carried out from both an ice camp and a research vessel in Baffin Bay, in the Canadian Arctic. This paper describes the sampling strategy and the dataset obtained from the research cruise, which took place aboard the Canadian Coast Guard ship (CCGS) Amundsen in late spring and early summer 2016. The sampling strategy was designed around the repetitive, perpendicular crossing of the marginal ice zone (MIZ), using not only ship-based station discrete sampling but also high-resolution measurements from autonomous platforms (Gliders, BGC-Argo floats …) and under-way monitoring systems. The dataset is available at https://doi.org/10.17882/86417 (Bruyant et al., 2022).

The Green Edge initiative was developed to investigate the processes controlling the primary productivity and fate of organic matter produced during the Arctic phytoplankton spring bloom (PSB) and to determine its role in the ecosystem. Two field campaigns were conducted in 2015 and 2016 at an ice camp located on landfast sea ice southeast of Qikiqtarjuaq Island in Baffin Bay (67.4797∘ N, 63.7895∘ W). During both expeditions, a large suite of physical, chemical and biological variables was measured beneath a consolidated sea-ice cover from the surface to the bottom (at 360 m depth) to better understand the factors driving the PSB. Key variables, such as conservative temperature, absolute salinity, radiance, irradiance, nutrient concentrations, chlorophyll a concentration, bacteria, phytoplankton and zooplankton abundance and taxonomy, and carbon stocks and fluxes were routinely measured at the ice camp. Meteorological and snow-relevant variables were also monitored. Here, we present the results of a joint effort to tidy and standardize the collected datasets, which will facilitate their reuse in other Arctic studies. The dataset is available at https://doi.org/10.17882/59892 (Massicotte et al., 2019a).

This article presents data regarding the Si biogeochemical cycle during two oceanographic cruises conducted in the tropical South Pacific (BIOSOPE and OUTPACE cruises) in 2005 and 2015. It involves the first Si stock measurements in this understudied region, encompassing various oceanic systems from New Caledonia to the Chilean upwelling between 8 and 34° S. Some of the lowest levels of biogenic silica standing stocks ever measured were found in this area, notably in the southern Pacific gyre, where Chlorophyll a concentrations are the most depleted worldwide. Integrated biogenic silica stocks are as low as 1.08±0.95 mmol m−2 and are the lowest stocks measured in the South Pacific. Size-fractionated biogenic silica concentrations revealed a non-negligible contribution of the pico-sized fraction (<2–3 µm) to biogenic silica standing stocks, representing 26% ± 12% of total biogenic silica during the OUTPACE cruise and 11% ± 9% during the BIOSOPE cruise. These results indicate significant accumulation in this size class, which was undocumented for 2005, but has since then been related to Si uptake by Synechococcus cells. Si uptake measurements carried out during BIOSOPE confirmed biological Si uptake by this size fraction. We further present diatoms community structure associated with the stock measurements for a global overview of the Si cycle in the tropical South Pacific.

Although the Southern Ocean is considered a high-nutrient, low-chlorophyll (HNLC) area, massive and recurrent blooms are observed over and downstream of the Kerguelen Plateau. This mosaic of blooms is triggered by a higher iron supply resulting from the interaction between the Antarctic Circumpolar Current and the local bathymetry. Net primary production, N uptake (NO3− and NH4+), and nitrification rates were measured at eight stations in austral spring 2011 (October–November) during the KEOPS 2 cruise in the Kerguelen Plateau area. Natural iron fertilization stimulated primary production, with mixed layer integrated net primary production and growth rates much higher in the fertilized areas (up to 315 mmol C m−2 d−1 and up to 0.31 d−1 respectively) compared to the HNLC reference site (12 mmol C m−2 d−1 and 0.06 d−1 respectively). Primary production was mainly sustained by nitrate uptake, with f ratios (corresponding to NO3−-uptake / (NO3−-uptake + NH4+-uptake)) lying at the upper end of the observations for the Southern Ocean (up to 0.9). We report high rates of nitrification (up to ~ 3 μmol N L−1 d−1, with ~ 90 % of them < 1 μmol N L−1 d−1) typically occurring below the euphotic zone, as classically observed in the global ocean. The specificity of the studied area is that at most of the stations, the euphotic layer was shallower than the mixed layer, implying that nitrifiers can efficiently compete with phytoplankton for the ammonium produced by remineralization at low-light intensities. Nitrate produced by nitrification in the mixed layer below the euphotic zone is easily supplied to the euphotic zone waters above, and nitrification sustained 70 ± 30 % of the nitrate uptake in the productive area above the Kerguelen Plateau. This complicates estimations of new production as potentially exportable production. We conclude that high productivity in deep mixing system stimulates the N cycle by increasing both assimilation and regeneration.

The biological composition of the material exported to a moored sediment trap located under the winter mixed layer of the naturally fertilized Kerguelen Plateau in the Southern Ocean was studied over an annual cycle. Despite iron availability in spring, the annual particulate organic carbon (POC) export (98.2 mmol m−2) at 289 m was low, but annual biogenic silica export was significant (114 mmol m−2). This feature was related to the abundance of empty diatom cells and the ratio of full to empty cells exerted a first-order control in BSi : POC export stoichiometry of the biological pump. Chaetoceros Hyalochaete spp. and Thalassiosira antarctica resting spores were responsible for more than 60% of the annual POC flux that occurred during two very short export events of < 14 days in spring–summer. Relatively low diatom fluxes were observed over the remainder of the year. Faecal pellet contribution to annual carbon flux was lower (34%) and reached its seasonal maximum in autumn and winter (> 80%). The seasonal progression of faecal pellet types revealed a clear transition from small spherical shapes (small copepods) in spring, to larger cylindrical and ellipsoid shapes in summer (euphausiids and large copepods) and finally to large tabular shapes (salps) in autumn and winter. We propose in this high-biomass, low-export (HBLE) environment that small but highly silicified and fast-sinking resting spores are able to bypass the intense grazing pressure and efficient carbon transfer to higher trophic levels that are responsible for the low fluxes observed the during the remainder of the year. More generally our study also provides a statistical framework linking the ecological succession of diatom and zooplankton communities to the seasonality of carbon and silicon export within an iron-fertilized bloom region in the Southern Ocean.

We examined phytoplankton community responses to natural iron fertilisation at 32 sites over and downstream from the Kerguelen Plateau in the Southern Ocean during the austral spring bloom in October–November 2011. The community structure was estimated from chemical and isotopic measurements (particulate organic carbon – POC; 13C-POC; particulate nitrogen – PN; 15N-PN; and biogenic silica – BSi) on size-fractionated samples from surface waters (300, 210, 50, 20, 5, and 1 μm fractions). Higher values of 13C-POC (vs. co-located 13C values for dissolved inorganic carbon – DIC) were taken as indicative of faster growth rates and higher values of 15N-PN (vs. co-located 15N-NO3 source values) as indicative of greater nitrate use (rather than ammonium use, i.e. higher f ratios). Community responses varied in relation to both regional circulation and the advance of the bloom. Iron-fertilised waters over the plateau developed dominance by very large diatoms (50–210 μm) with high BSi / POC ratios, high growth rates, and significant ammonium recycling (lower f ratios) as biomass built up. In contrast, downstream polar frontal waters with a similar or higher iron supply were dominated by smaller diatoms (20–50 μm) and exhibited greater ammonium recycling. Stations in a deep-water bathymetrically trapped recirculation south of the polar front with lower iron levels showed the large-cell dominance observed on the plateau but much less biomass. Comparison of these communities to surface water nitrate (and silicate) depletions as a proxy for export shows that the low-biomass recirculation feature had exported similar amounts of nitrogen to the high-biomass blooms over the plateau and north of the polar front. This suggests that early spring trophodynamic and export responses differed between regions with persistent low levels vs. intermittent high levels of iron fertilisation.

A massive diatom bloom is observed each year in the surface waters of the naturally Fe-fertilized Kerguelen Plateau (Southern Ocean). We measured biogenic silica production and dissolution fluxes (ρSi and ρDiss, respectively) in the mixed layer in the vicinity of the Kerguelen Plateau during austral spring 2011 (KEOPS-2 cruise). We compare results from a high-nutrient low-chlorophyll reference station and stations with different degrees of iron enrichment and bloom conditions. Above the plateau biogenic ρSi are among the highest reported so far in the Southern Ocean (up to 47.9 mmol m−2 d−1). Although significant (10.2 mmol m−2 d−1 on average), ρDiss were generally much lower than production rates. Uptake ratios (ρSi : ρC and ρSi : ρN) confirm that diatoms strongly dominate primary production in this area. At the bloom onset, decreasing dissolution-to-production ratios (D : P) indicate that the remineralization of silica could sustain most of the low silicon uptake and that the system progressively shifts toward a silica production regime which must be mainly supported by new source of silicic acid. Moreover, by comparing results from the two KEOPS expeditions (spring 2011 and summer 2005), we suggest that there is a seasonal evolution of the processes decoupling Si and N cycles in the area. Indeed, the consumption of H4SiO4 standing stocks occurs only during the growing stage of the bloom when strong net silica production is observed, contributing to higher H4SiO4 depletion relative to NO3−. Then, the decoupling of H4SiO4 and NO3− is mainly controlled by the more efficient nitrogen recycling relative to Si. Gross Si : N uptake ratios were higher in the Fe-rich regions compared to the high-nutrient low-chlorophyll (HNLC) area, likely due to different diatom communities. This suggests that the diatom responses to natural Fe fertilization are more complex than previously thought, and that natural iron fertilization over long timescales does not necessarily decrease Si : N uptake ratios as suggested by the silicic acid leakage hypothesis. Finally, we propose the first seasonal estimate of the Si biogeochemical budget above the Kerguelen Plateau based on direct measurements. This study points out that naturally iron-fertilized areas of the Southern Ocean could sustain very high regimes of biogenic silica production, similar to those observed in highly productive upwelling systems.

The particulate matter distribution and phytoplankton community structure of the iron-fertilized Kerguelen region were investigated in early austral spring (October–November 2011) during the KEOPS2 cruise. The iron-fertilized region was characterized by a complex mesoscale circulation resulting in a patchy distribution of particulate matter. Integrated concentrations over 200 m ranged from 72.2 to 317.7 mg m−2 for chlorophyll a 314 to 744 mmol m−2 for biogenic silica (BSi), 1106 to 2268 mmol m−2 for particulate organic carbon, 215 to 436 mmol m−2 for particulate organic nitrogen, and 29.3 to 39.0 mmol m−2 for particulate organic phosphorus. Three distinct high biomass areas were identified: the coastal waters of Kerguelen Islands, the easternmost part of the study area in the polar front zone, and the southeastern Kerguelen Plateau. As expected from previous artificial and natural iron-fertilization experiments, the iron-fertilized areas were characterized by the development of large diatoms revealed by BSi size–fractionation and high performance liquid chromatography (HPLC) pigment signatures, whereas the iron-limited reference area was associated with a low biomass dominated by a mixed (nanoflagellates and diatoms) phytoplankton assemblage. A major difference from most previous artificial iron fertilization studies was the observation of much higher Si : C, Si : N, and Si : P ratios (0.31 ± 0.16, 1.6 ± 0.7 and 20.5 ± 7.9, respectively) in the iron-fertilized areas compared to the iron-limited reference station (0.13, 1.1, and 5.8, respectively). A second difference is the patchy response of the elemental composition of phytoplankton communities to large scale natural iron fertilization. Comparison to the previous KEOPS1 cruise also allowed to address the seasonal dynamics of phytoplankton bloom over the southeastern plateau. From particulate organic carbon (POC), particulate organic nitrogen (PON), and BSi evolutions, we showed that the elemental composition of the particulate matter also varies at the seasonal scale. This temporal evolution followed changes of the phytoplankton community structure as well as major changes in the nutrient stocks progressively leading to silicic acid exhaustion at the end of the productive season. Our observations suggest that the specific response of phytoplankton communities under natural iron fertilization is much more diverse than what has been regularly observed in artificial iron fertilization experiments and that the elemental composition of the bulk particulate matter reflects phytoplankton taxonomic structure rather than being a direct consequence of iron availability.

Phytoplankton identification and abundance data are now commonly feeding plankton distribution databases worldwide. This study is a first attempt to compile the largest possible body of data available from different databases as well as from individual published or unpublished datasets regarding diatom distribution in the world ocean. The data obtained originate from time series studies as well as spatial studies. This effort is supported by the Marine Ecosystem Model Inter-Comparison Project (MAREMIP), which aims at building consistent datasets for the main plankton functional types (PFTs) in order to help validate biogeochemical ocean models by using carbon (C) biomass derived from abundance data. In this study we collected over 293 000 individual geo-referenced data points with diatom abundances from bottle and net sampling. Sampling site distribution was not homogeneous, with 58% of data in the Atlantic, 20% in the Arctic, 12% in the Pacific, 8% in the Indian and 1% in the Southern Ocean. A total of 136 different genera and 607 different species were identified after spell checking and name correction. Only a small fraction of these data were also documented for biovolumes and an even smaller fraction was converted to C biomass. As it is virtually impossible to reconstruct everyone's method for biovolume calculation, which is usually not indicated in the datasets, we decided to undertake the effort to document, for every distinct species, the minimum and maximum cell dimensions, and to convert all the available abundance data into biovolumes and C biomass using a single standardized method. Statistical correction of the database was also adopted to exclude potential outliers and suspicious data points. The final database contains 90 648 data points with converted C biomass. Diatom C biomass calculated from cell sizes spans over eight orders of magnitude. The mean diatom biomass for individual locations, dates and depths is 141.19 μg C l−1, while the median value is 11.16 μg C l−1. Regarding biomass distribution, 19% of data are in the range 0–1 μg C l−1, 29% in the range 1–10 μg C l−1, 31% in the range 10–100 μg C l−1, 18% in the range 100–1000 μg C l−1, and only 3% > 1000 μg C l−1. Interestingly, less than 50 species contributed to > 90% of global biomass, among which centric species were dominant. Thus, placing significant efforts on cell size measurements, process studies and C quota calculations of these species should considerably improve biomass estimates in the upcoming years. A first-order estimate of the diatom biomass for the global ocean ranges from 444 to 582 Tg C, which converts to 3 to 4 Tmol Si and to an average Si biomass turnover rate of 0.15 to 0.19 d−1. Link to the dataset: doi:10.1594/PANGAEA.777384.

The silicon biogeochemical cycle has been studied in the Mediterranean Sea during late summer/early autumn 1999 and summer 2008. The distribution of nutrients, particulate carbon and silicon, fucoxanthin (Fuco), and total chlorophyll-a (TChl-a) were investigated along an eastward gradient of oligotrophy during two cruises (PROSOPE and BOUM) encompassing the entire Mediterranean Sea during the stratified period. At both seasons, surface waters were depleted in nutrients and the nutriclines gradually deepened towards the East, the phosphacline being the deepest in the easternmost Levantine basin. Following the nutriclines, parallel deep maxima of biogenic silica (DSM), fucoxanthin (DFM) and TChl-a (DCM) were evidenced during both seasons with maximal concentrations of 0.45 μmol L−1 for BSi, 0.26 μg L−1 for Fuco, and 1.70 μg L−1 for TChl-a, all measured during summer. Contrary to the DCM which was a persistent feature in the Mediterranean Sea, the DSM and DFMs were observed in discrete areas of the Alboran Sea, the Algero-Provencal basin, the Ionian sea and the Levantine basin, indicating that diatoms were able to grow at depth and dominate the DCM under specific conditions. Diatom assemblages were dominated by Chaetoceros spp., Leptocylindrus spp., Pseudonitzschia spp. and the association between large centric diatoms (Hemiaulus hauckii and Rhizosolenia styliformis) and the cyanobacterium Richelia intracellularis was observed at nearly all sites. The diatom's ability to grow at depth is commonly observed in other oligotrophic regions and could play a major role in ecosystem productivity and carbon export to depth. Contrary to the common view that Si and siliceous phytoplankton are not major components of the Mediterranean biogeochemistry, we suggest here that diatoms, by persisting at depth during the stratified period, could contribute to a large part of the marine primary production as observed in other oligotrophic areas.

A one-year survey of the sedimenting particulate material was conducted at the SOFi site located on the edge of the continental slope in the Gulf of Lions (Northwestern Mediterranean) between September 1999 and December 2000. The main focus of this paper was to establish an annual budget of biogenic silica export, to describe the seasonnal pattern of Si fluxes in relation to the organic and mineral C fluxes and to quantify the Si/C decoupling processes during sedimentation. The pattern of the downward flux of BSi was partly typical with increased daily sedimentation rates during spring followed by smaller peaks during summer. However, an unusual sedimentation event was recorded in the bottom trap in February before the spring thermal stratification occurred, and represented 37% of the annual BSi mass flux. The total annual BSi flux at the SOFi site was estimated from the bottom trap and amounted to 86.8 mmol Si m-2 y-1, while the organic carbon flux amounted to 154.9 mmol C m-2 y-1. The overall efficiency of C export to depth compared to surface primary production budgets was low (3%) and similar to values found for oligotrophic regions of the Eastern Mediterranean. On the other hand, we documented a very high efficiency of the Si export at the SOFi site, with 62% of the Si produced in the surface layer exported to the bottom trap. The excellent correlation between BSi and particulate organic C (POC) in the bottom trap further indicates that the bulk of the organic matter is transferred to depth in association with diatoms. However, the main fraction of the C produced by phytoplankton is lost in the water column through oxidation or metabolisation, while biogenic silica is more efficiently transferred to depth. This strong Si/C decoupling with depth is likely to be the cause of the spring depletion of silicic acid over the entire water column which was observed by mid-April in a companion study (Leblanc et al., 2003) and may cause a severe Si limitation of the diatom spring bloom in this oligotrophic region of the Gulf of Lions.

Afin d'analyser la réponse de l'écosystème « Rade de Brest » à l'enrichissement du milieu, un suivi du cycle des concentrations en sels nutritifs et en chlorophylle a été réalisé en 1993 et ces résultats ont été comparés à des études antérieures. Bien que les apports de nitrates aient doublé en 20 ans, occasionnant une diminution des ratios Si/N, les stocks phytoplanctoniques n'ont pas augmenté durant cette période. L'analyse des données ainsi que la modélisation écologique ont permis de mettre en évidence plusieurs facteurs de régulation du milieu, contribuant à expliquer la relative stabilité de cet écosystème malgré l'augmentation des apports de nitrate.

Diatoms are one of the major primary producers in the ocean, responsible annually for ~20% of photosynthetically fixed CO 2 on Earth. In oceanic models, they are typically represented as large (>20 µm) microphytoplankton. However, many diatoms belong to the nanophytoplankton (2-20 µm) and a few species even overlap with the picoplanktonic size-class (<2 µm). Due to their minute size and difficulty of detection they are poorly characterized. Here we describe a massive spring bloom of the smallest known diatom (Minidiscus) in the northwestern Mediterranean Sea. Analysis of Tara Oceans data, together with literature review, reveal a general oversight of the significance of these small diatoms at the global scale. We further evidence that they can reach the seafloor at high sinking rates, implying the need to revise our classical binary vision of pico- and nanoplanktonic cells fueling the microbial loop, while only microphytoplankton sustain secondary trophic levels and carbon export.

In the naturally iron-fertilized surface waters of the northern Kerguelen Plateau region, the early spring diatom community composition and contribution to plankton carbon biomass were investigated and compared with the high nutrient, low chlorophyll (HNLC) surrounding waters. The large iron-induced blooms were dominated by small diatom species belonging to the genera Chaetoceros (Hyalochaete) and Thalassiosira, which rapidly responded to the onset of favorable light-conditions in the meander of the Polar Front. In comparison, the iron-limited HNLC area was typically characterized by autotrophic nanoeukaryote-dominated communities and by larger and more heavily silicified diatom species (e.g. Fragilariopsis spp.). Our results support the hypothesis that diatoms are valuable vectors of carbon export to depth in naturally iron-fertilized systems of the Southern Ocean. Furthermore, our results corroborate observations of the exported diatom assemblage from a sediment trap deployed in the iron-fertilized area, whereby the dominant Chaetoceros (Hyalochaete) cells were less efficiently exported than the less abundant, yet heavily silicified, cells of Thalassionema nitzschioides and Fragilariopsis kerguelensis Our observations emphasize the strong influence of species-specific diatom cell properties combined with trophic interactions on matter export efficiency, and illustrate the tight link between the specific composition of phytoplankton communities and the biogeochemical properties characterizing the study area., (© FEMS 2016. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

The availability of iron limits primary productivity and the associated uptake of carbon over large areas of the ocean. Iron thus plays an important role in the carbon cycle, and changes in its supply to the surface ocean may have had a significant effect on atmospheric carbon dioxide concentrations over glacial-interglacial cycles. To date, the role of iron in carbon cycling has largely been assessed using short-term iron-addition experiments. It is difficult, however, to reliably assess the magnitude of carbon export to the ocean interior using such methods, and the short observational periods preclude extrapolation of the results to longer timescales. Here we report observations of a phytoplankton bloom induced by natural iron fertilization--an approach that offers the opportunity to overcome some of the limitations of short-term experiments. We found that a large phytoplankton bloom over the Kerguelen plateau in the Southern Ocean was sustained by the supply of iron and major nutrients to surface waters from iron-rich deep water below. The efficiency of fertilization, defined as the ratio of the carbon export to the amount of iron supplied, was at least ten times higher than previous estimates from short-term blooms induced by iron-addition experiments. This result sheds new light on the effect of long-term fertilization by iron and macronutrients on carbon sequestration, suggesting that changes in iron supply from below--as invoked in some palaeoclimatic and future climate change scenarios--may have a more significant effect on atmospheric carbon dioxide concentrations than previously thought.

The net inputs of silicic acid (dissolved silica) to the world ocean have been revised to 6.1 +/- 2.0 teramoles of silicon per year (1 teramole = 10(12) moles). The major contribution (about 80 percent) comes from rivers, whose world average silicic acid concentration is 150 micromolar. These inputs are reasonably balanced by the net ouputs of biogenic silica of 7.1 +/- 1.8 teramoles of silicon per year in modern marine sediments. The gross production of biogenic silica (the transformation of dissolved silicate to particulate skeletal material) in surface waters was estimated to be 240 +/- 40 teramoles of silicon per year, and the preservation ratio (opal accumulation in sediment/gross production in surface waters) averages 3 percent. In the world ocean the residence time of silicon, relative to total biological uptake in surface waters, is about 400 years.