Articles | Volume 12, issue 4
Research article 26 Feb 2015
Research article | 26 Feb 2015
Vertical partitioning of phosphate uptake among picoplankton groups in the low Pi Mediterranean Sea
A. Talarmin et al.
No articles found.
Biogeosciences, 16, 1673–1673,
Pascale Bouruet-Aubertot, Yannis Cuypers, Andrea Doglioli, Mathieu Caffin, Christophe Yohia, Alain de Verneil, Anne Petrenko, Dominique Lefèvre, Hervé Le Goff, Gilles Rougier, Marc Picheral, and Thierry Moutin
Biogeosciences, 15, 7485–7504,Short summary
The OUTPACE cruise took place between New Caledonia and French Polynesia. The main purpose was to understand how micro-organisms can survive in a very poor environment. One main source of nutrients is at depth, below the euphotic layer where micro-organisms live. The purpose of the turbulence measurements was to determine to which extent turbulence may
upliftnutrients into the euphotic layer. The origin of the turbulence that was found contrasted along the transect was also determined.
Audrey Gimenez, Melika Baklouti, Thibaut Wagener, and Thierry Moutin
Biogeosciences, 15, 6573–6589,Short summary
During the OUTPACE cruise conducted in the oligotrophic to ultra-oligotrophic region of the western tropical South Pacific, two contrasted regions were sampled in terms of N2 fixation rates, primary production rates and nutrient availability. The aim of this work was to investigate the role of N2 fixation in the differences observed between the two contrasted areas by comparing two simulations only differing by the presence or not of N2 fixers using a 1-D biogeochemical–physical coupled model.
Thibaut Wagener, Nicolas Metzl, Mathieu Caffin, Jonathan Fin, Sandra Helias Nunige, Dominique Lefevre, Claire Lo Monaco, Gilles Rougier, and Thierry Moutin
Biogeosciences, 15, 5221–5236,Short summary
The western tropical South Pacific was sampled along a longitudinal 4000 km transect (OUTPACE cruise) for the measurement of carbonate parameters (total alkalinity and total inorganic carbon) between the Melanesian Archipelago and the western part of the South Pacific gyre. This paper reports this new dataset and derived properties. We also estimate anthropogenic carbon distribution in the water column using the TrOCA method.
Thierry Moutin, Thibaut Wagener, Mathieu Caffin, Alain Fumenia, Audrey Gimenez, Melika Baklouti, Pascale Bouruet-Aubertot, Mireille Pujo-Pay, Karine Leblanc, Dominique Lefevre, Sandra Helias Nunige, Nathalie Leblond, Olivier Grosso, and Alain de Verneil
Biogeosciences, 15, 2961–2989,Short summary
Surface waters of the western tropical South Pacific were sampled along a longitudinal 4000 km transect during the stratified period between the Melanesian Archipelago and the western part of the South Pacific gyre. We found a significant biological carbon pump sustained almost exclusively by N2 fixation and essentially controlled by phosphate availability in the iron-rich Melanesian Archipelago waters which appears to be a net sink for atmospheric CO2 while the gyre is in a quasi-steady state.
Angela N. Knapp, Kelly M. McCabe, Olivier Grosso, Nathalie Leblond, Thierry Moutin, and Sophie Bonnet
Biogeosciences, 15, 2619–2628,Short summary
The spatial distribution of biological N2 fixation fluxes to the ocean remains poorly constrained. Here we use nitrogen isotope budgets to identify significant N2 fixation inputs to the western tropical South Pacific (WTSP), where N2 fixation supports > 50 % of export production at stations proximal to iron sources. The significant N2 fixation inputs in the WTSP may offset nitrogen loss in the oxygen-deficient zones of the eastern tropical South Pacific.
France Van Wambeke, Audrey Gimenez, Solange Duhamel, Cécile Dupouy, Dominique Lefevre, Mireille Pujo-Pay, and Thierry Moutin
Biogeosciences, 15, 2669–2689,Short summary
The western tropical South Pacific Ocean has recently been shown to be a hotspot for biological nitrogen fixation. In this study, we examined the horizontal and vertical distribution of heterotrophic prokaryotic production alongside photosynthetic rates, nitrogen fixation rates and phosphate turnover times across the western tropical South Pacific Ocean, in order to relate these fluxes to bottom–up controls (related to nitrogen, phosphate and labile C availability).
Mathieu Caffin, Thierry Moutin, Rachel Ann Foster, Pascale Bouruet-Aubertot, Andrea Michelangelo Doglioli, Hugo Berthelot, Cécile Guieu, Olivier Grosso, Sandra Helias-Nunige, Nathalie Leblond, Audrey Gimenez, Anne Alexandra Petrenko, Alain de Verneil, and Sophie Bonnet
Biogeosciences, 15, 2565–2585,Short summary
We performed N budgets to assess the role of N2 fixation on production and export in the western tropical South Pacific Ocean. We deployed a combination of techniques including high-sensitivity measurements of N input and sediment traps deployment. We demonstrated that N2 fixation was the major source of new N before atmospheric deposition and upward nitrate fluxes. It contributed significantly to organic matter export, indicating a high efficiency of this region to export carbon.
Alain de Verneil, Louise Rousselet, Andrea M. Doglioli, Anne A. Petrenko, Christophe Maes, Pascale Bouruet-Aubertot, and Thierry Moutin
Biogeosciences, 15, 2125–2147,Short summary
Oceanographic campaigns to measure biogeochemical processes popularly deploy drifters with onboard incubations to stay in a single body of water. Here, we aggregate physical data taken during such a cruise, OUTPACE, to independently test in a new approach whether the drifter really stayed in what can be considered a single biological or chemical environment. This study concludes that future campaigns would benefit from similar data collection and analysis to validate their sampling strategy.
Alain Fumenia, Thierry Moutin, Sophie Bonnet, Mar Benavides, Anne Petrenko, Sandra Helias Nunige, and Christophe Maes
Revised manuscript not acceptedShort summary
The Melanesian archipelago waters between 160° E and 170° W are characterized by a significant N2 fixation rates and an excess of particulate organic nitrogen compared to the canonical ratio of Redfield and a positive N*. We hypothesize that the southern branch of the subtropical gyre is probably the main vector of excess nitrogen transport in the thermocline waters showing an influence of nitrogen fixation occurring in the western tropical in a large part of the South Pacific.
Alain de Verneil, Louise Rousselet, Andrea M. Doglioli, Anne A. Petrenko, and Thierry Moutin
Biogeosciences, 14, 3471–3486,Short summary
A surface summer plankton bloom in the western tropical South Pacific was sampled during the Oligotrophy to UlTra-oligotrophy PACific Experiment (OUTPACE) cruise. We characterize the bloom's properties and the circulation responsible for its evolution. Nitrogen fixation helped sustain the bloom, and larger-scale flows, rather than the smaller ones, explain its movements. Future studies of blooms in this region can make use of these findings to track the horizontal export of plankton production.
Thierry Moutin, Andrea Michelangelo Doglioli, Alain de Verneil, and Sophie Bonnet
Biogeosciences, 14, 3207–3220,Short summary
The overall goal of OUTPACE was to obtain a successful representation of the interactions between planktonic organisms and the cycle of biogenic elements in the western tropical South Pacific Ocean across trophic and N2 fixation gradients. The international OUTPACE cruise took place between 18 February and 3 April 2015 aboard the RV L’Atalante and involved 60 scientists. The transect covered ~4 000 km from the western part of the Melanesian archipelago to the western boundary of the gyre.
Audrey Gimenez, Melika Baklouti, Sophie Bonnet, and Thierry Moutin
Biogeosciences, 13, 5103–5120,Short summary
In the context of the VAHINE mesocosm experiment in the Nouméa lagoon (New Caledonia), a 1-D vertical biogeochemical mechanistic model was used together with the in situ experiment to complement our comprehension of the planktonic ecosystem dynamics and the main biogeochemical carbon, nitrogen and phosphate fluxes. The model also showed the fate of fixed N2 by providing, over time, the proportion of diazotroph-derived nitrogen (DDN) in each compartment (mineral and organic) of the model.
Angela N. Knapp, Sarah E. Fawcett, Alfredo Martínez-Garcia, Nathalie Leblond, Thierry Moutin, and Sophie Bonnet
Biogeosciences, 13, 4645–4657,Short summary
The goal of this manuscript was to track the fate of newly fixed nitrogen (N) in large volume mesocosms in the coastal waters of New Caledonia. We used a N isotope ("δ15N") budget and found a shift in the δ15N of sinking particulate N over the 23-day experiment, indicating that nitrate supported export production at the beginning of the experiment, but that nitrogen fixation supported export at the end. We infer that nitrogen fixation supported export production by a release of dissolved N.
Ilana Berman-Frank, Dina Spungin, Eyal Rahav, France Van Wambeke, Kendra Turk-Kubo, and Thierry Moutin
Biogeosciences, 13, 3793–3805,Short summary
In the marine environment, sticky sugar-containing gels, termed transparent exopolymeric particles (TEP), are produced from biological sources and physical and chemical processes. These compounds are essential vectors enhancing downward flow of organic matter and its storage at depth. Spatial and temporal dynamics of TEPs were followed for 23 days during the VAHINE mesocosm experiment that investigated the fate of nitrogen and carbon derived from organisms fixing atmospheric N2 (diazotrophs).
France Van Wambeke, Ulrike Pfreundt, Aude Barani, Hugo Berthelot, Thierry Moutin, Martine Rodier, Wolfgang R. Hess, and Sophie Bonnet
Biogeosciences, 13, 3187–3202,Short summary
The phytoplankton is at the base of the plankton food web in large parts of oceanic "deserts" such as the South Pacific Ocean, where nitrogen sources limit activity. Mesocosms were fertilized with phosphorus to stimulate diazotrophy (atmospheric N2 fixation). Mostly diazotroph-derived nitrogen fuelled the heterotrophic bacterial community through indirect processes generating dissolved organic matter and detritus, such as mortality, lysis and grazing of both diazotrophs and non-diazotrophs.
Sophie Bonnet, Thierry Moutin, Martine Rodier, Jean-Michel Grisoni, Francis Louis, Eric Folcher, Bertrand Bourgeois, Jean-Michel Boré, and Armelle Renaud
Biogeosciences, 13, 2803–2814,Short summary
e main goal of the VAHINE project was to study the fate of N2 fixation in the ocean. Three large-volume (~ 50 m3) mesocosms were deployed in a tropical oligotrophic ecosystem (the New Caledonia lagoon, south-eastern Pacific). This introductory paper describes the scientific objectives of the project in detail as well as the implementation plan: the mesocosm description and deployment, the selection of the study site, and the logistical and sampling strategy.
A. Guyennon, M. Baklouti, F. Diaz, J. Palmieri, J. Beuvier, C. Lebaupin-Brossier, T. Arsouze, K. Béranger, J.-C. Dutay, and T. Moutin
Biogeosciences, 12, 7025–7046,Short summary
Dissolved organic carbon (DOC) has already been identified as a potentially significant source of carbon export in the Mediterranean Sea, though in situ export estimations are scarce. This work provides a thorough analysis at basin scale of carbon export with the coupled model NEMO-MED12/Eco3M-MED model. The seasonality and the processes of particulate and dissolved carbon production are also investigated. DOC export appears to be dominant in most regions, especially in the eastern basin.
H. Berthelot, T. Moutin, S. L'Helguen, K. Leblanc, S. Hélias, O. Grosso, N. Leblond, B. Charrière, and S. Bonnet
Biogeosciences, 12, 4099–4112,
Related subject area
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acidity variability and extremes under increasing atmospheric CO2Can ocean community production and respiration be determined by measuring high-frequency oxygen profiles from autonomous floats?Oxygen budget for the north-western Mediterranean deep convection regionAssessing the value of biogeochemical Argo profiles versus ocean color observations for biogeochemical model optimization in the Gulf of MexicoReviews and syntheses: The biogeochemical cycle of silicon in the modern oceanThe Southern Annular Mode (SAM) influences phytoplankton communities in the seasonal ice zone of the Southern OceanLinking intrinsic and apparent relationships between phytoplankton and environmental forcings using machine learning ‐ What are the challenges?Contrasted release of insoluble elements (Fe, Al, REE, Th, Pa) after dust deposition in seawater: a tank experiment approachProfiling float observation of thermohaline staircases in the western Mediterranean Sea and impact on nutrient fluxesOcean carbonate 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PacificQuantitative mapping and predictive modeling of Mn nodules' distribution from hydroacoustic and optical AUV data linked by random forests machine learningArtificial radionuclides in neon flying squid from the northwestern Pacific in 2011 following the Fukushima accident
Fuminori Hashihama, Hiroaki Saito, Taketoshi Kodama, Saori Yasui-Tamura, Jota Kanda, Iwao Tanita, Hiroshi Ogawa, E. Malcolm S. Woodward, Philip W. Boyd, and Ken Furuya
Biogeosciences, 18, 897–915,Short summary
We investigated the nutrient assimilation characteristics of deep-water-induced phytoplankton blooms across the subtropical North and South Pacific Ocean. Nutrient drawdown ratios of dissolved inorganic nitrogen to phosphate were anomalously low in the western North Pacific, likely due to the high phosphate uptake capability of low-phosphate-adapted phytoplankton. The anomalous phosphate uptake might influence the maintenance of chronic phosphate depletion in the western North Pacific.
Florian Ricour, Arthur Capet, Fabrizio D'Ortenzio, Bruno Delille, and Marilaure Grégoire
Biogeosciences, 18, 755–774,Short summary
This paper addresses the phenology of the deep chlorophyll maximum (DCM) in the Black Sea (BS). We show that the DCM forms in March at a density level set by the winter mixed layer. It maintains this location until June, suggesting an influence of the DCM on light and nutrient profiles rather than mere adaptation to external factors. In summer, the DCM concentrates ~55 % of the chlorophyll in a 10 m layer at ~35 m depth and should be considered a major feature of the BS phytoplankton dynamics.
Robyn E. Tuerena, Joanne Hopkins, Raja S. Ganeshram, Louisa Norman, Camille de la Vega, Rachel Jeffreys, and Claire Mahaffey
Biogeosciences, 18, 637–653,Short summary
The Barents Sea is a rapidly changing shallow sea within the Arctic. Here, nitrate, an essential nutrient, is fully consumed by algae in surface waters during summer months. Nitrate is efficiently regenerated in the Barents Sea, and there is no evidence for nitrogen loss from the sediments by denitrification, which is prevalent on other Arctic shelves. This suggests that nitrogen availability in the Barents Sea is largely determined by the supply of nutrients in water masses from the Atlantic.
Biogeosciences, 18, 509–534,Short summary
Biogeochemical-Argo floats are starting to routinely measure ocean chlorophyll, nutrients, oxygen, and pH. This study generated synthetic observations representing two potential Biogeochemical-Argo observing system designs and created a data assimilation scheme to combine them with an ocean model. The proposed system of 1000 floats brought clear benefits to model results, with additional floats giving further benefit. Existing satellite ocean colour observations gave complementary information.
Mark Hague and Marcello Vichi
Biogeosciences, 18, 25–38,Short summary
This paper examines the question of what causes the rapid spring growth of microscopic marine algae (phytoplankton) in the ice-covered ocean surrounding Antarctica. One prominent hypothesis proposes that the melting of sea ice is the primary cause, while our results suggest that this is only part of the explanation. In particular, we show that phytoplankton are able to start growing before the sea ice melts appreciably, much earlier than previously thought.
Arthur Capet, Luc Vandenbulcke, and Marilaure Grégoire
Biogeosciences, 17, 6507–6525,Short summary
The Black Sea is 2000 m deep, but, due to limited ventilation, only about the upper 100 m contains enough oxygen to support marine life such as fish. This oxygenation depth has been shown to be decreasing (1955–2019). Here, we evidence that atmospheric warming induced a clear shift in an important ventilation mechanism. We highlight the impact of this shift on oxygenation. There are important implications for marine life and carbon and nutrient cycling if this new ventilation regime persists.
Tim Rixen, Greg Cowie, Birgit Gaye, Joaquim Goes, Helga do Rosário Gomes, Raleigh R. Hood, Zouhair Lachkar, Henrike Schmidt, Joachim Segschneider, and Arvind Singh
Biogeosciences, 17, 6051–6080,Short summary
The northern Indian Ocean hosts an extensive oxygen minimum zone (OMZ), which intensified due to human-induced global changes. This includes the occurrence of anoxic events on the Indian shelf and affects benthic ecosystems and the pelagic ecosystem structure in the Arabian Sea. Consequences for biogeochemical cycles are unknown, which, in addition to the poor representation of mesoscale features, reduces the reliability of predictions of the future OMZ development in the northern Indian Ocean.
Marion Lagarde, Nolwenn Lemaitre, Hélène Planquette, Mélanie Grenier, Moustafa Belhadj, Pascale Lherminier, and Catherine Jeandel
Biogeosciences, 17, 5539–5561,
Natacha Le Grix, Jakob Zscheischler, Charlotte Laufkötter, Cécile S. Rousseaux, and Thomas L. Frölicher
Revised manuscript accepted for BGShort summary
Marine ecosystems could suffer severe damage from the co-occurrence of a marine heatwave with extremely low chlorophyll concentration. Here, we provide a first assessment of compound marine heatwave and low chlorophyll events in the global ocean from 1998 to 2018. We reveal hotspots of these compound events in the equatorial Pacific and in the Arabian Sea, show that they mostly occur in summer at high latitudes, and that their frequency is modulated by large-scale modes of climate variability.
Randelle M. Bundy, Alessandro Tagliabue, Nicholas J. Hawco, Peter L. Morton, Benjamin S. Twining, Mariko Hatta, Abigail E. Noble, Mattias R. Cape, Seth G. John, Jay T. Cullen, and Mak A. Saito
Biogeosciences, 17, 4745–4767,Short summary
Cobalt (Co) is an essential nutrient for ocean microbes and is scarce in most areas of the ocean. This study measured Co concentrations in the Arctic Ocean for the first time and found that Co levels are extremely high in the surface waters of the Canadian Arctic. Although the Co primarily originates from the shelf, the high concentrations persist throughout the central Arctic. Co in the Arctic appears to be increasing over time and might be a source of Co to the North Atlantic.
Friedrich A. Burger, Jasmin G. John, and Thomas L. Frölicher
Biogeosciences, 17, 4633–4662,Short summary
Ensemble simulations of an Earth system model reveal that ocean acidity extremes have increased in the past few decades and are projected to increase further in terms of frequency, intensity, duration, and volume extent. The increase is not only caused by the long-term ocean acidification due to the uptake of anthropogenic CO2, but also due to changes in short-term variability. The increase in ocean acidity extremes may enhance the risk of detrimental impacts on marine organisms.
Christopher Gordon, Katja Fennel, Clark Richards, Lynn K. Shay, and Jodi K. Brewster
Biogeosciences, 17, 4119–4134,Short summary
We describe a method for correcting errors in oxygen optode measurements on autonomous platforms in the ocean. The errors result from the relatively slow response time of the sensor. The correction method includes an in situ determination of the effective response time and requires the time stamps of the individual measurements. It is highly relevant for the BGC-Argo program and also applicable to gliders. We also explore if diurnal changes in oxygen can be obtained from profiling floats.
Caroline Ulses, Claude Estournel, Marine Fourrier, Laurent Coppola, Fayçal Kessouri, Dominique Lefèvre, and Patrick Marsaleix
Revised manuscript accepted for BGShort summary
We analyze the seasonal cycle of O2 and estimate an annual O2 budget in the north-western Mediterranean deep convection region, using a numerical model. We show that this region acts as a large sink of atmospheric O2 and as a major source of O2 for the western Mediterranean Sea. The decrease in the deep convection intensity predicted in recent projections may have important consequences on the overall uptake of O2 in the Mediterranean Sea and on the O2 exchanges with the Atlantic Ocean.
Bin Wang, Katja Fennel, Liuqian Yu, and Christopher Gordon
Biogeosciences, 17, 4059–4074,Short summary
We assess trade-offs between different types of biological observations, specifically satellite ocean color and BGC-Argo profiles and the benefits of combining both for optimizing a biogeochemical model of the Gulf of Mexico. Using all available observations leads to significant improvements in observed and unobserved variables (including primary production and C export). Our results highlight the significant benefits of BGC-Argo measurements for biogeochemical model optimization and validation.
Paul J. Tréguer, Jill N. Sutton, Mark Brzezinski, Matthew A. Charette, Timothy Devries, Stephanie Dutkiewicz, Claudia Ehlert, Jon Hawkings, Aude Leynaert, Su Mei Liu, Natalia Llopis Monferrer, María López-Acosta, Manuel Maldonado, Shaily Rahman, Lihua Ran, and Olivier Rouxel
Revised manuscript accepted for BGShort summary
Silicon is the second-most abundant element of the Earth's crust. In this review, we show that silicon inputs and outputs, to and from the world ocean, are 57 % and 18 % higher, respectively, than previous estimates. These changes are significant, modifying factors such as the geochemical residence time of silicon, which is now about 8000 years and two times faster than previously assumed. We also update the total biogenic silica pelagic production and provide an estimate for sponge production.
Bruce L. Greaves, Andrew T. Davidson, Alexander D. Fraser, John P. McKinlay, Andrew Martin, Andrew McMinn, and Simon W. Wright
Biogeosciences, 17, 3815–3835,Short summary
We observed that variation in the Southern Annular Mode (SAM) over 11 years showed a relationship with the species composition of hard-shelled phytoplankton in the seasonal ice zone (SIZ) of the Southern Ocean. Phytoplankton in the SIZ are productive during the southern spring and summer when the area is ice-free, with production feeding most Antarctic life. The SAM is known to be increasing with climate change, and changes in phytoplankton in the SIZ may have implications for higher life forms.
Christopher Holder and Anand Gnanadesikan
Revised manuscript accepted for BGShort summary
A frequent challenge for marine ecologists is linking small-scale relationships found in a lab to broader relationships observed on large scales in the environment. We investigated whether machine learning (ML) could help connect these small and large-scale relationships. ML can recover the small-scale relationships when the small and large-scale relationships operate on similar time and spatial scales, but this breaks down when the variability in the observations is lost due to time-averaging.
Matthieu Roy-Barman, Lorna Folio, Eric Douville, Nathalie Leblond, Fréderic Gazeau, Matthieu Bressac, Thibaut Wagener, Céline Ridame, Karine Desboeufs, and Cécile Guieu
Revised manuscript under review for BGShort summary
The release of insoluble elements such as Aluminum (Al), Iron (Fe), Rare Earth Elements (REE), Thorium (Th) and Protactinium (Pa) when Saharan dust fall over the Mediterranean Sea was studied during tank experiments, under present and future climate conditions. Each element exhibited different dissolution kinetics and dissolution fractions (always lower than a few percent). Changes in temperature and/or pH under greenhouse conditions lead to a lower Th release and a higher light REE release.
Vincent Taillandier, Louis Prieur, Fabrizio D'Ortenzio, Maurizio Ribera d'Alcalà, and Elvira Pulido-Villena
Biogeosciences, 17, 3343–3366,Short summary
This study addresses the role played by vertical diffusion in the nutrient enrichment of the Levantine intermediate waters, a process particularly relevant inside thermohaline staircases. Thanks to a high profiling frequency over a 4-year period, BGC-Argo float observations reveal the temporal continuity of the layering patterns encountered during the cruise PEACETIME and their impact on vertical and lateral transfers of nitrate between the deep reservoir and the surface productive zone.
Coraline Leseurre, Claire Lo Monaco, Gilles Reverdin, Nicolas Metzl, Jonathan Fin, Solveig Olafsdottir, and Virginie Racapé
Biogeosciences, 17, 2553–2577,Short summary
In this study, we investigate the evolution of CO2 uptake and ocean acidification in the North Atlantic Subpolar surface water. Our results show an important reduction in the capacity of the ocean to absorb CO2 from the atmosphere (1993–2007), due to a rapid increase in the fCO2 and associated with a rapid decrease in pH. Conversely, data obtained during the last decade (2008–2017) show a stagnation of fCO2 (increasing the ocean sink for CO2) and pH.
Antonio Tovar-Sánchez, Araceli Rodríguez-Romero, Anja Engel, Birthe Zäncker, Franck Fu, Emilio Marañón, María Pérez-Lorenzo, Matthieu Bressac, Thibaut Wagener, Sylvain Triquet, Guillaume Siour, Karine Desboeufs, and Cécile Guieu
Biogeosciences, 17, 2349–2364,Short summary
Residence times of particulate metals derived from aerosol deposition in the Sea Surface Microlayer of the Mediterranean Sea ranged from a couple of minutes (e.g., for Fe) to a few hours (e.g., for Cu). Microbial activity seems to play an important role in in this process and in the concentration and distribution of metals between diferent water layers.
Pieter Demuynck, Toby Tyrrell, Alberto Naveira Garabato, Mark Christopher Moore, and Adrian Peter Martin
Biogeosciences, 17, 2289–2314,Short summary
The availability of macronutrients N and Si is of key importance to sustain life in the Southern Ocean. N and Si are available in abundance at the southern boundary of the Southern Ocean due to constant supply from the deep ocean. In the more northern regions of the Southern Ocean, a decline in macronutrient concentration is noticed, especially strong for Si rather than N. This paper uses a simplified biogeochemical model to investigate processes responsible for this decline in concentration.
Martine Lizotte, Maurice Levasseur, Virginie Galindo, Margaux Gourdal, Michel Gosselin, Jean-Éric Tremblay, Marjolaine Blais, Joannie Charette, and Rachel Hussherr
Biogeosciences, 17, 1557–1581,Short summary
This study brings further support to the premise that the prevalence of younger and thinner icescapes over older and thicker ones in the Canadian High Arctic favors the early development of under-ice microorganisms as well as their production of the climate-relevant gas dimethylsulfide (DMS). Given the rapid rate of climate-driven changes in Arctic sea ice, our results suggest implications for the timing and magnitude of DMS pulses in the Arctic, with ramifications for climate forecasting.
Mark J. Hopwood, Nicolas Sanchez, Despo Polyviou, Øystein Leiknes, Julián Alberto Gallego-Urrea, Eric P. Achterberg, Murat V. Ardelan, Javier Aristegui, Lennart Bach, Sengul Besiktepe, Yohann Heriot, Ioanna Kalantzi, Tuba Terbıyık Kurt, Ioulia Santi, Tatiana M. Tsagaraki, and David Turner
Biogeosciences, 17, 1309–1326,Short summary
Hydrogen peroxide, H2O2, is formed naturally in sunlight-exposed water by photochemistry. At high concentrations it is undesirable to biological cells because it is a stressor. Here, across a range of incubation experiments in diverse marine environments (Gran Canaria, the Mediterranean, Patagonia and Svalbard), we determine that two factors consistently affect the H2O2 concentrations irrespective of geographical location: bacteria abundance and experiment design.
Neil J. Wyatt, Angela Milne, Eric P. Achterberg, Thomas J. Browning, Heather A. Bouman, E. Malcolm S. Woodward, and Maeve C. Lohan
Revised manuscript accepted for BGShort summary
Using data collected during two expeditions to the South Atlantic ocean, we investigated how the interaction between external sources and biological activity influenced the availability of the trace metals zinc and cobalt. This is important as both metals play essential roles in the metabolism and growth of phytoplankton and thus influence primary productivity of the oceans. We found seasonal changes in both processes that helped explain upper ocean trace metal cycling.
Manon Tonnard, Hélène Planquette, Andrew R. Bowie, Pier van der Merwe, Morgane Gallinari, Floriane Desprez de Gésincourt, Yoan Germain, Arthur Gourain, Marion Benetti, Gilles Reverdin, Paul Tréguer, Julia Boutorh, Marie Cheize, François Lacan, Jan-Lukas Menzel Barraqueta, Leonardo Pereira-Contreira, Rachel Shelley, Pascale Lherminier, and Géraldine Sarthou
Biogeosciences, 17, 917–943,Short summary
We investigated the spatial distribution of dissolved Fe during spring 2014, in order to understand the processes influencing the biogeochemical cycle in the North Atlantic. Our results highlighted elevated Fe close to riverine inputs at the Iberian Margin and glacial inputs at the Newfoundland and Greenland margins. Atmospheric deposition appeared to be a minor source of Fe. Convection was an important source of Fe in the Irminger Sea, which was depleted in Fe relative to nitrate.
Carolin R. Löscher, Wiebke Mohr, Hermann W. Bange, and Donald E. Canfield
Biogeosciences, 17, 851–864,Short summary
Oxygen minimum zones (OMZs) are ocean areas severely depleted in oxygen as a result of physical, chemical, and biological processes. Biologically, organic material is produced in the sea surface and exported to deeper waters, where it respires. In the Bay of Bengal (BoB), an OMZ is present, but there are traces of oxygen left. Our study now suggests that this is because one key process, nitrogen fixation, is absent in the BoB, thus preventing primary production and consecutive respiration.
Lothar Stramma, Sunke Schmidtko, Steven J. Bograd, Tsuneo Ono, Tetjana Ross, Daisuke Sasano, and Frank A. Whitney
Biogeosciences, 17, 813–831,Short summary
The influence of climate signals in the Pacific, especially the Pacific Decadal Oscillation and the North Pacific Gyre Oscillation, as well as El Niño–La Niña and an 18.6-year nodal tidal cycle on oxygen and nutrient trends is investigated. At different locations in the Pacific Ocean different climate signals dominate. Hence, not only trends related to warming but also the influence of climate signals need to be investigated to understand oxygen and nutrient changes in the ocean.
Marie-Hélène Radenac, Julien Jouanno, Christine Carine Tchamabi, Mesmin Awo, Bernard Bourlès, Sabine Arnault, and Olivier Aumont
Biogeosciences, 17, 529–545,Short summary
Satellite data and a remarkable set of in situ measurements show a main bloom of microscopic seaweed, the phytoplankton, in summer and a secondary bloom in December in the central equatorial Atlantic. They are driven by a strong vertical supply of nitrate in May–July and a shorter and moderate supply in November. In between, transport of low-nitrate water from the west explains most nitrate losses in the sunlit layer. Horizontal eddy-induced processes also contribute to seasonal nitrate removal.
Andrés S. Rigual Hernández, Thomas W. Trull, Scott D. Nodder, José A. Flores, Helen Bostock, Fátima Abrantes, Ruth S. Eriksen, Francisco J. Sierro, Diana M. Davies, Anne-Marie Ballegeer, Miguel A. Fuertes, and Lisa C. Northcote
Biogeosciences, 17, 245–263,Short summary
Coccolithophores account for a major fraction of the carbonate produced in the world's oceans. However, their contribution in the subantarctic Southern Ocean remains undocumented. We quantitatively partition calcium carbonate fluxes amongst coccolithophore species in the Australian–New Zealand sector of the Southern Ocean. We provide new insights into the importance of species other than Emiliania huxleyi in the carbon cycle and assess their possible response to projected environmental change.
Susana Agustí, Jeffrey W. Krause, Israel A. Marquez, Paul Wassmann, Svein Kristiansen, and Carlos M. Duarte
Biogeosciences, 17, 35–45,Short summary
We found that 24 % of the total diatoms community in the Arctic water column (450 m depth) was located below the photic layer. Healthy diatom communities in active spring–bloom stages remained in the photic layer. Dying diatom communities exported a large fraction of the biomass to the aphotic zone, fuelling carbon sequestration and benthic ecosystems in the Arctic. The results of the study conform to a conceptual model where diatoms grow during the bloom until silicic acid stocks are depleted.
Xinwei Wang, Feixue Fu, Pingping Qu, Joshua D. Kling, Haibo Jiang, Yahui Gao, and David A. Hutchins
Biogeosciences, 16, 4393–4409,Short summary
In this study, we examine the responses of E. huxleyi to a future warmer and more thermally variable ocean. Elevated temperatures and thermal variation have negative effects on growth rate and physiology that are especially pronounced at high temperatures, but high-frequency thermal variation may reduce the risk of extreme high-temperature events. These findings have potentially large implications for ocean productivity and marine biogeochemical cycles under a future changing climate.
Federico Baltar and Gerhard J. Herndl
Biogeosciences, 16, 3793–3799,Short summary
Around half of the global primary production (PP) is produced in the ocean. Here we quantified how much oceanic PP estimates would increase if we included the dark DIC fixation rates (which are usually excluded in the carbon-14 method) into the PP estimation. We found that the inclusion of dark DIC fixation would increase PP estimates by 5–22 %. This represents ca. 1.2 to 11 Pg C yr−1 of newly synthesized organic carbon available for the marine food web.
Renaud Person, Olivier Aumont, Gurvan Madec, Martin Vancoppenolle, Laurent Bopp, and Nacho Merino
Biogeosciences, 16, 3583–3603,Short summary
The Antarctic Ice Sheet is considered a possibly important but largely overlooked source of iron (Fe). Here we explore its fertilization capacity by evaluating the response of marine biogeochemistry to Fe release from icebergs and ice shelves in a global ocean model. Large regional impacts are simulated, leading to only modest primary production and carbon export increases at the scale of the Southern Ocean. Large uncertainties are due to low observational constraints on modeling choices.
Robyn E. Tuerena, Raja S. Ganeshram, Matthew P. Humphreys, Thomas J. Browning, Heather Bouman, and Alexander P. Piotrowski
Biogeosciences, 16, 3621–3635,Short summary
The carbon isotopes in algae can be used to predict food sources and environmental change. We explore how dissolved carbon is taken up by algae in the South Atlantic Ocean and how this affects their carbon isotope signature. We find that cell size controls isotope fractionation. We use our results to investigate how climate change may impact the carbon isotopes in algae. We suggest a shift to smaller algae in this region would decrease the carbon isotope ratio at the base of the food web.
Daniela Niemeyer, Iris Kriest, and Andreas Oschlies
Biogeosciences, 16, 3095–3111,Short summary
Recent studies suggest spatial variations of the marine particle flux length scale. Using a global biogeochemical ocean model, we investigate whether changes in particle size and size-dependent sinking can explain this variation. We address uncertainties by varying aggregate properties and circulation. Both aspects have an impact on the representation of nutrients, oxygen and oxygen minimum zones. The formation and sinking of large aggregates in productive areas lead to deeper flux penetration.
Jamie D. Wilson, Stephen Barker, Neil R. Edwards, Philip B. Holden, and Andy Ridgwell
Biogeosciences, 16, 2923–2936,Short summary
The remains of plankton rain down from the surface ocean to the deep ocean, acting to store CO2 in the deep ocean. We used a model of biology and ocean circulation to explore the importance of this process in different regions of the ocean. The amount of CO2 stored in the deep ocean is most sensitive to changes in the Southern Ocean. As plankton in the Southern Ocean are likely those most impacted by future climate change, the amount of CO2 they store in the deep ocean could also be affected.
Natalie C. Harms, Niko Lahajnar, Birgit Gaye, Tim Rixen, Kirstin Dähnke, Markus Ankele, Ulrich Schwarz-Schampera, and Kay-Christian Emeis
Biogeosciences, 16, 2715–2732,Short summary
The Indian Ocean subtropical gyre is a large oligotrophic area that is likely to adjust to continued warming by increasing stratification, reduced nutrient supply and decreasing biological production. In this study, we investigated concentrations of nutrients and stable isotopes of nitrate. We determine the lateral influence of water masses entering the gyre from the northern Indian Ocean and from the Southern Ocean and quantify the input of nitrogen by N2 fixation into the surface layer.
Yingxu Wu, Mathis P. Hain, Matthew P. Humphreys, Sue Hartman, and Toby Tyrrell
Biogeosciences, 16, 2661–2681,Short summary
This study takes advantage of the GLODAPv2 database to investigate the processes driving the surface ocean dissolved inorganic carbon distribution, with the focus on its latitudinal gradient between the polar oceans and the low-latitude oceans. Based on our quantitative study, we find that temperature-driven CO2 gas exchange and high-latitude upwelling of DIC- and TA-rich deep waters are the two major drivers, with the importance of the latter not having been previously realized.
Qixing Ji, Mark A. Altabet, Hermann W. Bange, Michelle I. Graco, Xiao Ma, Damian L. Arévalo-Martínez, and Damian S. Grundle
Biogeosciences, 16, 2079–2093,Short summary
A strong El Niño event occurred in the Peruvian coastal region in 2015–2016, during which higher sea surface temperatures co-occurred with significantly lower sea-to-air fluxes of nitrous oxide, an important greenhouse gas and ozone depletion agent. Stratified water column during El Niño retained a larger amount of nitrous oxide that was produced via multiple microbial pathways; and intense nitrous oxide effluxes could occur when normal upwelling is resumed after El Niño.
Ulrike Löptien and Heiner Dietze
Biogeosciences, 16, 1865–1881,Short summary
Anthropogenic greenhouse gas emissions trigger complex climate feedbacks. Output form Earth system models provides a basis for related political decision-making. One challenge is to arrive at reliable model parameter estimates for the ocean biogeochemistry module. We illustrate pitfalls through which flaws in the ocean module are masked by wrongly tuning the biogeochemistry and discuss ensuing uncertainties in climate projections.
Arthur Gourain, Hélène Planquette, Marie Cheize, Nolwenn Lemaitre, Jan-Lukas Menzel Barraqueta, Rachel Shelley, Pascale Lherminier, and Géraldine Sarthou
Biogeosciences, 16, 1563–1582,Short summary
The GEOVIDE cruise (May–June 2014, R/V Pourquoi Pas?) aimed to provide a better understanding of trace metal biogeochemical cycles in the North Atlantic. As particles play a key role in the global biogeochemical cycle of trace elements in the ocean, we discuss the distribution of particulate iron (PFe). Lithogenic sources appear to dominate the PFe cycle through margin and benthic inputs.
Jan-Lukas Menzel Barraqueta, Jessica K. Klar, Martha Gledhill, Christian Schlosser, Rachel Shelley, Hélène F. Planquette, Bernhard Wenzel, Geraldine Sarthou, and Eric P. Achterberg
Biogeosciences, 16, 1525–1542,Short summary
We used surface water dissolved aluminium concentrations collected in four different GEOTRACES cruises to determine atmospheric deposition fluxes to the ocean. We calculate atmospheric deposition fluxes for largely under-sampled regions of the Atlantic Ocean and thus provide new constraints for models of atmospheric deposition. The use of the MADCOW model is of major importance as dissolved aluminium is analysed within the GEOTRACES project at high spatial resolution.
Karin F. Kvale, Katherine E. Turner, Angela Landolfi, and Katrin J. Meissner
Biogeosciences, 16, 1019–1034,Short summary
Drivers motivating the evolution of calcifying phytoplankton are poorly understood. We explore differences in global ocean chemistry with and without calcifiers during rapid climate changes. We find the presence of phytoplankton calcifiers stabilizes the volume of low oxygen regions and consequently stabilizes the concentration of nitrate, which is an important nutrient required for photosynthesis. By stabilizing nitrate concentrations, calcifiers improve their growth conditions.
Debany Fonseca-Batista, Xuefeng Li, Virginie Riou, Valérie Michotey, Florian Deman, François Fripiat, Sophie Guasco, Natacha Brion, Nolwenn Lemaitre, Manon Tonnard, Morgane Gallinari, Hélène Planquette, Frédéric Planchon, Géraldine Sarthou, Marc Elskens, Julie LaRoche, Lei Chou, and Frank Dehairs
Biogeosciences, 16, 999–1017,Short summary
Dinitrogen fixation and primary production were investigated using stable isotope incubation experiments along two transects off the Western Iberian Margin in May 2014 close to the end of the phytoplankton spring bloom. We observed substantial N2 fixation activities (up to 1533 µmol N m-2 d-1) associated with a predominance of unicellular cyanobacteria and non-cyanobacterial diazotrophs, which seemed to be promoted by the presence of bloom-derived organic matter and excess phosphorus.
Ludivine Conte, Sophie Szopa, Roland Séférian, and Laurent Bopp
Biogeosciences, 16, 881–902,Short summary
The ocean is a source of atmospheric carbon monoxide, a key component for the oxidizing capacity of the atmosphere. We use a global ocean biogeochemistry model to dynamically assess the oceanic CO budget and its emission to the atmosphere at the global scale. The total emissions of CO to the atmosphere are 4.0 Tg C yr−1. The oceanic CO emission maps produced are relevant for use by atmospheric chemical models, especially to study the oxidizing capacity of the atmosphere above the remote ocean.
Yi Tang, Nolwenn Lemaitre, Maxi Castrillejo, Montserrat Roca-Martí, Pere Masqué, and Gillian Stewart
Biogeosciences, 16, 309–327,Short summary
Oceanographers try to understand the ocean’s role in the global carbon cycle. Trace levels of natural radionuclides can inform this connection and their half-lives provide an estimate of the timing of processes. We used the 210Po and 210Pb pair to examine the export of carbon from the surface ocean to depth along the GEOVIDE GEOTRACES cruise track. We found that the flux was regionally variable, that upwelling was an important regional factor, and that both large and small particles drove flux.
Christos Panagiotopoulos, Mireille Pujo-Pay, Mar Benavides, France Van Wambeke, and Richard Sempéré
Biogeosciences, 16, 105–116,
Iason-Zois Gazis, Timm Schoening, Evangelos Alevizos, and Jens Greinert
Biogeosciences, 15, 7347–7377,Short summary
The use of high-resolution hydroacoustic and optic data acquired by an autonomous underwater vehicle can give us detailed sea bottom topography and valuable information regarding manganese nodules' spatial distribution. Moreover, the combined use of these data sets with a random forest machine learning model can extend this spatial prediction beyond the areas with available photos, providing researchers with a new mapping tool for further investigation and links with other data.
Wen Yu, Mathew P. Johansen, Jianhua He, Wu Men, and Longshan Lin
Biogeosciences, 15, 7235–7242,Short summary
To better understand the impact of the Fukushima accident on commercial marine species, neon flying squid samples obtained from the NW Pacific in Nov 2011 were analyzed for a range of radionuclides. Elevated levels of Cs-134 and Ag-110m from the Fukushima accident were found in the samples, with an extremely high concentration ratio for Ag-110m. However, the radiological dose for squid living in the study area, and for human consumers of these squid, was far below the recommended dose limits.
Agusti, S.: Viability and niche segregation of Prochlorococcus and Synechococcus cells across the Central Atlantic Ocean, Aquat. Microb. Ecol., 36, 53–59, 2004.
Azam, F., Fenchel, T., Field, J. G., Gray, J. S., Meyer-Reil, L. A., and Thingstad, F.: The Ecological Role of Water-Column Microbes in the Sea, Mar. Ecol.-Prog. Ser., 10, 257–263, 1983.
Bertilsson, S., Berglund, O., Karl, D. M., and Chisholm, S. W.: Elemental Composition of Marine Prochlorococcus and Synechococcus: implications for the Ecological Stoichiometry of the Sea, Limnol. Oceanogr., 48, 1721–1731, 2003.
Björkman, K. and Karl, D. M.: Bioavailability of inorganic and organic phosphorus compounds to natural assemblages of microorganisms in Hawaiian coastal waters, Mar. Ecol.-Prog. Ser., 111, 265–273, 1994.
Björkman, K., Duhamel, S., and Karl, D. M.: Microbial Group Specific Uptake Kinetics of Inorganic Phosphate and Adenosine-5`-Triphosphate (ATP) in the North Pacific Subtropical Gyre, Frontiers in Microbiology, 3, 189, https://doi.org/10.3389/fmicb.2012.00189, 2012.
Casey, J. R., Lomas, M. W., Mandecki, J., and Walker, D. E.: Prochlorococcus contributes to new production in the Sargasso Sea deep chlorophyll maximum Geophys. Res. Lett., 34, L10604, https://doi.org/10.1029/2006GL028725, 2007.
Casey, J. R., Lomas, M. W., Michelou, V. K., Dyhrman, S. T., Orchard, E. D., Ammerman, J. W., and Sylvan, J. B.: Phytoplankton taxon-specific orthophosphate (Pi) and ATP utilization in the western subtropical North Atlantic, Aquat. Microb. Ecol., 58, 31–44, https://doi.org/10.3354/ame01348, 2009.
Christaki, U., Van Wambeke, F., and Dolan, J. R.: Nanoflagellates (mixotrophs, heterotrophs and autotrophs) in the oligotrophic eastern Mediterranean: standing stocks, bacterivory and relationships with bacterial production, Mar. Ecol.-Prog. Ser., 181, 297–307, 1999.
Church, M. J., Ducklow, H., and Karl, D. M.: Multiyear increases in dissolved organic matter inventories at Station ALOHA in the North Pacific Subtropical Gyre, Limnol. Oceanogr., 47, 1–10, 2002.
Cotner, J. B. and Biddanda, B. A.: Small Players, Large Role: Microbial Influence on Biogeochemical Processes in Pelagic Aquatic Ecosystems, Ecosystems, 5, 105–121, 2002.
Crombet, Y., Leblanc, K., Quéguiner, B., Moutin, T., Rimmelin, P., Ras, J., Claustre, H., Leblond, N., Oriol, L., and Pujo-Pay, M.: Deep silicon maxima in the stratified oligotrophic Mediterranean Sea, Biogeosciences, 8, 459–475, https://doi.org/10.5194/bg-8-459-2011, 2011.
Currie, D. J. and Kalff, J.: A comparison of the abilities of freshwater algae and bacteria to acquire and retain phosphorus, Limnol. Oceanogr., 29, 298–310, 1984.
Currie, D. J., Bentzen, E., and Kalff, J.: Does algal-bacterial phosphorus partitioning vary among lakes? a comparative study of orthophosphate uptake and alkaline phosphatase activity in freshwater, Can. J. Fish. Aquat. Sci., 43, 311–318, https://doi.org/10.1139/f86-040, 1986.
Dao, M. H.: Reassessment of the cell surface area limitation to nutrient uptake in phytoplankton, Mar. Ecol. Prog. Ser., 489, 87–92, https://doi.org/10.3354/meps10434, 2013.
Drakare, S.: Competition between Picoplanktonic Cyanobacteria and Heterotrophic Bacteria along Crossed Gradients of Glucose and Phosphate, Microbial Ecol., 44, 327–335, https://doi.org/10.1007/s00248-002-1013-4, 2002.
Duhamel, S., Björkman, K. M., and Karl, D. M.: Light dependence of phosphorus uptake by microorganisms in the subtropical North and South Pacific Ocean, Aquat. Microb. Ecol., 67, 225–238, https://doi.org/10.3354/ame01593, 2012.
Flaten, G. A. F., Skjoldal, E. F., Krom, M. D., Law, C. S., Mantoura, R. F. C., Paraskevi, P., Psarra, S., Tanaka, T., Tselepides, A., Woodward, E. M. S., Zohary, T., and Thingstad, T. F.: Studies of the microbial P-cycle during a Lagrangian phosphate-addition experiment in the Eastern Mediterranean, Deep Sea Res. Pt. II, 52, 2928–2943, 2005.
Fu, F. X., Zhang, Y., Feng, Y., and Hutchins, D. A.: Phosphate and ATP uptake and growth kinetics in axenic cultures of the cyanobacterium Synechococcus CCMP 1334, Eur. J. Phycol., 41, 15–28, 2006.
Hartmann, M., Grob, C., Scanlan, D. J., Martin, A. P., Burkill, P. H., and Zubkov, M. V.: Comparison of phosphate uptake rates by the smallest plastidic and aplastidic protists in the North Atlantic subtropical gyre, FEMS Microbiol. Ecol., 78, 327–335, https://doi.org/10.1111/j.1574-6941.2011.01160.x, 2011.
Ho, T.-Y., Quigg, A., Finkel, Z. V., Milligan, A. J., Wyman, K., Falkowski, P. G., and Morel, F. M. M.: The elemental composition of some marine phytoplankton, J. Phycol., 39, 1145–1159, https://doi.org/10.1111/j.0022-3646.2003.03-090.x, 2003.
Karl, D. M.: Microbially Mediated Transformations of Phosphorus in the Sea: New Views of an Old Cycle, Annual Review of Marine Science, 6, 279–337, https://doi.org/10.1146/annurev-marine-010213-135046, 2014.
Karl, D. M. and Tien, G.: MAGIC: A sensitive and precise method for measuring dissolved phosphorus in aquatic environments, Limnol. Oceanogr., 37, 105–116, 1992.
Karl, D. M., Bidigare, R. R., and Letelier, R. M.: Long-term changes in plankton community structure and productivity in the North Pacific Subtropical Gyre: The domain shift hypothesis, Deep Sea Res. Pt. II, 48, 1449–1470, https://doi.org/10.1016/S0967-0645(00)00149-1, 2001.
Kashtan, N., Roggensack, S., Rodrigue, S., Thompson, J. W., Biller, S. J., Coe, A., DIng, H., Marttinen, P., Malmstrom, R. R., Stocker, R., Follows, M. J., Stepanauskas, R., and Chisholm, S. W.: Single-cell genomics reveals hundreds of coexisting subpopulations in wild Prochlorococcus, Science, 344, 416–420, https://doi.org/10.1126/science.1248575, 2014.
Knauss, H. J. and Porter, J. W.: The absorption of inorganic ions by Chlorella pyrenoidosa, Plant Physiol., 29, 229–234, 1954.
Krom, M. D., Woodward, E. M. S., Herut, B., Kress, N., Carbo, P., Mantoura, R. F. C., Spyres, G., Thingstad, T. F., Wassmann, P., Wexels-Riser, C., Kitidis, V., Law, C. S., and Zodiatis, G.: Nutrient cycling in the south east Levantine basin of the eastern Mediterranean: Results from a phosphorus starved system, Deep Sea Res. Pt. II, 52, 2879–2896, 2005.
Labry, C., Herbland, A., and Delmas, D.: The role of phosphorus on planktonic production of the Gironde plume waters in the Bay of Biscay, J. Plankton Res., 24, 97–117, https://doi.org/10.1093/plankt/24.2.97, 2002.
Lomas, M. W., Burke, A. L., Lomas, D. A., Bell, D. W., Shen, C., Dyhrman, S. T., and Ammerman, J. W.: Sargasso Sea phosphorus biogeochemistry: an important role for dissolved organic phosphorus (DOP), Biogeosciences, 7, 695–710, https://doi.org/10.5194/bg-7-695-2010, 2010.
Lomas, M. W., Bonachela, J. A., Levin, S. A., and Martiny, A. C.: Impact of ocean phytoplankton diversity on phosphate uptake, P. Natl. A. Sci., 111, 17540–17545, https://doi.org/10.1073/pnas.1420760111, 2014.
Marie, D., Simon, N., Guillou, L., Partensky, F., and Vaulot, D.: DNA/RNA Analysis of Phytoplankton by Flow Cytometry UNIT 11.12, Current Protocols in Cytometry, 2000.
Marie, D., Shi, X. L., Rigaut-Jalabert, F., and Vaulot, D.: Use of flow cytometric sorting to better assess the diversity of small photosynthetic eukaryotes in the English Channel, FEMS Microbiol. Ecol., 72, 165–178, https://doi.org/10.1111/j.1574-6941.2010.00842.x, 2010.
Martínez, A., Osburne, M. S., Sharma, A. K., DeLong, E. F., and Chisholm, S. W.: Phosphite utilization by the marine picocyanobacterium Prochlorococcus MIT9301, Environ. Microbiol., 14, 1363–1377, https://doi.org/10.1111/j.1462-2920.2011.02612.x, 2012.
McLaughlin, K., Sohm, J. A., Cutter, G. A., Lomas, M. W., and Paytan, A.: Phosphorus cycling in the Sargasso Sea: Investigation using the oxygen isotopic composition of phosphate, enzyme-labeled fluorescence, and turnover times, Global Biogeochem. Cy., 27, 375–387, https://doi.org/10.1002/gbc.20037, 2013.
Mella-Flores, D., Mazard, S., Humily, F., Partensky, F., Mahé, F., Bariat, L., Courties, C., Marie, D., Ras, J., Mauriac, R., Jeanthon, C., Mahdi Bendif, E., Ostrowski, M., Scanlan, D. J., and Garczarek, L.: Is the distribution of Prochlorococcus and Synechococcus ecotypes in the Mediterranean Sea affected by global warming?, Biogeosciences, 8, 2785–2804, https://doi.org/10.5194/bg-8-2785-2011, 2011.
Michelou, V. K., Lomas, M. W., and Kirchman, D. L.: Phosphate and adenosine-5'-triphosphate uptake by cyanobacteria and heterotrophic bacteria in the Sargasso Sea, Limnol. Oceanogr., 56, 323–332, 2011.
Moutin, T. and Prieur, L.: Influence of anticyclonic eddies on the Biogeochemistry from the Oligotrophic to the Ultraoligotrophic Mediterranean (BOUM cruise), Biogeosciences, 9, 3827–3855, https://doi.org/10.5194/bg-9-3827-2012, 2012.
Moutin, T., Thingstad, T. F., Van Wambeke, F., Marie, D., Slawyk, G., Raimbault, P., and Claustre, H.: Does Competition for Nanomolar Phosphate Supply Explain the Predominance of the Cyanobacterium Synechococcus?, Limnol. Oceanogr., 47, 1562–1567, 2002.
Nyholm, N.: Kinetics of phosphate limited algal growth, Biotechnol. Bioeng., 19, 467–492, https://doi.org/10.1002/bit.260190404, 1977.
Paerl, H. and Merkel, S.: The effects of particles on phosphorus assimilation in attached vs. free floating microorganisms, Arch. Hydrobiol, 93, 125–134, 1982.
Paffenhöfer, G. A., Sherr, B. F., and Sherr, E. B.: From small scales to the big picture: persistence mechanisms of planktonic grazers in the oligotrophic ocean, Mar. Ecol., 28, 243–253, 2007.
Rhee, G. Y.: A continuous culture study of phosphate uptake, growth rate and polyphosphate in Scenedesmus sp., J. Phycol., 9, 495–506, https://doi.org/10.1111/j.1529-8817.1973.tb04126.x, 1973.
Rimmelin, P. and Moutin, T.: Re-examination of the MAGIC method to determine low orthophosphate concentration in seawater, Anal. Chim. Acta, 548, 174–182, 2005.
Sebastián, M., Pitta, P., González, J. M., Thingstad, T. F., and Gasol, J. M.: Bacterioplankton groups involved in the uptake of phosphate and dissolved organic phosphorus in a mesocosm experiment with P-starved Mediterranean waters, Environ. Microbiol., 14, 2334–2347, https://doi.org/10.1111/j.1462-2920.2012.02772.x, 2012.
Simon, M.: Specific uptake rates of amino acids by attached and free-living bacteria in a mesotrophic lake, Appl. Environ. Microb., 49, 1254–1259, 1985.
Talarmin, A., Van Wambeke, F., Catala, P., Courties, C., and Lebaron, P.: Flow cytometric assessment of specific leucine incorporation in the open Mediterranean, Biogeosciences, 8, 253–265, https://doi.org/10.5194/bg-8-253-2011, 2011a.
Talarmin, A., Van Wambeke, F., Duhamel, S., Catala, P., Moutin, T., and Lebaron, P.: Improved methodology to measure taxon-specific phosphate uptake in live and unfiltered samples, Limnol. Oceanogr.-Meth., 9, 443–453, https://doi.org/10.4319/lom.2011.9.443, 2011b.
Tanaka, T., Rassoulzadegan, F., and Thingstad, T. F.: Measurements of phosphate affinity constants and phosphorus release rates from the microbial food web in Villefranche Bay, northwestern Mediterranean, Limnol. Oceanogr., 48, 1150–1160, 2003.
Tanaka, T., Thingstad, T. F., Christaki, U., Colombet, J., Cornet-Barthaux, V., Courties, C., Grattepanche, J.-D., Lagaria, A., Nedoma, J., Oriol, L., Psarra, S., Pujo-Pay, M., and Van Wambeke, F.: Lack of P-limitation of phytoplankton and heterotrophic prokaryotes in surface waters of three anticyclonic eddies in the stratified Mediterranean Sea, Biogeosciences, 8, 525–538, https://doi.org/10.5194/bg-8-525-2011, 2011.
Ternon, E., Guieu, C., Ridame, C., L'Helguen, S., and Catala, P.: Longitudinal variability of the biogeochemical role of Mediterranean aerosols in the Mediterranean Sea, Biogeosciences, 8, 1067–1080, https://doi.org/10.5194/bg-8-1067-2011, 2011.
Thingstad, T. F., Skjoldal, E. F., and Bohne, R. A.: Phosphorus cycling and algal-bacterial competition in Sandsfjord, western Norway, Mar. Ecol. Prog. Ser., 99, 239–259, 1993.
Thingstad, T. F., Zweifel, U. L., and Rassoulzadegan, F.: P limitation of heterotrophic bacteria and phytoplankton in the northwest Mediterranean, Limnol. Oceanogr., 43, 88–94, 1998.
Webb, M. R.: A continuous spectrophotometric assay for inorganic phosphate and for measuring phosphate release kinetics in biological systems, P. Natl. Acad. Sci. USA, 89, 4884–4887, 1992.
Wu, J., Sunda, W., Boyle, E. A., and Karl, D. M.: Phosphate Depletion in the Western North Atlantic Ocean, Science, 289, 759–762, https://doi.org/10.1126/science.289.5480.759, 2000.
Zubkov, M. V., Mary, I., Woodward, E. M. S., Warwick, P. E., Fuchs, B. M., Scanlan, D. J., and Burkill, P. H.: Microbial control of phosphate in the nutrient-depleted North Atlantic subtropical gyre, Environ. Microbiol., 9, 2079–2089, 2007.
Along a transect in the warm nutrient-depleted Mediterranean Sea, we found that microorganisms shared phosphate resources by using different uptake strategies. Heterotrophic prokaryotes dominated phosphate uptake in the upper layers while cyanobacteria dominated uptake fluxes around the deep chlorophyll maximum depth. Synechococcus seemed well equipped to respond to pulses of phosphate, whereas Prochlorococcus and heterotrophs were more adapted to very low concentrations.
Along a transect in the warm nutrient-depleted Mediterranean Sea, we found that microorganisms...