Distribution of inorganic and organic nutrients in the South Pacific Ocean – evidence for long-term accumulation of organic matter in nitrogen-depleted waters

International audienceThe BIOSOPE cruise of the RV Atalante was devoted to study the biogeochemical properties in the South Pacific between the Marquesas Islands (141° W–8° S) and the Chilean upwelling (73° W–34° S). The 8000 km cruise had the opportunity to encounter different trophic situations, and especially strong oligotrophic conditions in the Central South Pacific Gyre (SPG, between 123° W and 101° W). In this isolated region, nitrate was undetectable between surface and 160–180 m, while regenerated nitrogen (nitrite and ammonium) only revealed some traces (-1), even in the subsurface maximum. Integrated nitrate over the photic layer, which reached 165 m, was close to zero. In spite of this severe nitrogen-depletion, phosphate was always present at significant concentrations (˜0.1 µmoles l-1), while silicate maintained at low but classical oceanic levels (˜1 µmoles l-1). In contrast, the Marquesas region (MAR) at west and Chilean upwelling (UPW) at east were characterized by large nutrient contents one hundred to one thousand fold higher than in the SPG. Distribution of surface chlorophyll concentration reflected this gradient of nitrate availability. The lowest value (0.023 nmoles l-1) was measured in the centre of the SPG, where integrated chlorophyll over the photic layer was very weak (˜10 mg m-2), since a great part (up to 50%) of the deep chlorophyll maximum (DCM) was located below the 1% light. But, because of the relative high concentration encountered in the DCM (0.2 µg l-1), chlorophyll a content over the photic layer varied much less (by a factor 2 to 5) than the nitrate content. In contrast to chlorophyll a, integrated content of particulate organic matter (POM) remained more or less constant along the investigated area (500 mmoles m-2, 60 mmoles m-2 and 3.5 mmoles m-2 for particulate organic carbon, particulate organic nitrogen and particulate organic phosphorus, respectively), except in the upwelling where values were two fold higher. Extensive comparison has shown that glass fiber GF/F filters efficiency collected particulate chlorophyll, while a significant fraction of POM (up to 50%) passed trough this filter and was retained by 0.2 µm Teflon membrane. The most striking feature was the large accumulation of dissolved organic matter (DOM) in the SPG relative to surrounding waters, especially dissolved organic carbon (DOC) where concentrations were at levels rarely measured in oceanic waters (>100 µmoles l-1). Due to this large pool of DOM over the whole photic layer of the SPG, integrated values followed an opposite geographical pattern than this of inorganic nutrients with a large accumulation within the centre of the SPG. While suspended particulate matter in the mixed layer had C/N ratio largely conform to Redfield stoichiometry (C/N˜6.6), marked deviations were observed in this excess DOM (C/N˜16 to 23). The existence of C-rich dissolved organic matter is recognized as a feature typical of oligotrophic waters, requiring the over consumption of carbon. Thus, in spite of strong nitrate-depletion leading to low chlorophyll biomass, the closed ecosystem of the SPG can produce a large amount of carbon. The implications of this finding are discussed, the conclusion being that, due to the lack of seasonal vertical mixing and weak lateral advection, the dissolved organic carbon biologically produced can be accumulated and stored in the photic layer for a very long period

Two High-Nutrient Low-Chlorophyll phytoplankton assemblages: the tropical central Pacific and the offshore Perú-Chile Current

International audienceThe phytoplankton (>15 µm) composition and abundance was investigated along a ~8000 km transect between the Marquesas Islands Archipelago and the Chilean coasts off Concepción. In the southern limit of the central Equatorial Pacific (at 8° S, 141° W), in High-Nutrient Low-Chlorophyll (HNLC) warm waters, the microphytoplankton assemblage was dominated by the lightly silicified diatoms Pseudo-nitzschia delicatissima and Rhizosolenia bergonii. The morphology of these species, a small pennate diatom that exhibited a tendency to form "ball of needles" clusters and large centric diatom (>500 µm long), are interpreted as two anti-grazing strategies in an environment dominated by small micrograzers. Surprisingly, this a priori typical HNLC phytoplankton assemblage was also found in the temperate offshore waters of the Perú-Chile Current between 2000 and 600 km off Chile. This observation suggests that a common set of environmental factors (obviously other than temperature and salinity) are responsible for the establishment and maintaining of this distinctive phytoplankton in these geographically and hydrologically distant regions. Both regions are characterized by a surface nitrate-silicic acid ratio ranging from 1–3. Occasionally Rhizosolenia bergonii showed frustules anomalously fragmented, likely the result of extreme weakly silicified phytoplankton. We suggest that silicon deficiency may be responsible of the occurrence of HNLC phytoplankton assemblage in the tropical central Pacific as well as the offshore Perú-Chile Current during the austral summer

Dissolved organic matter uptake by <i>Trichodesmium </i>in the Southwest Pacific

International audienceThe globally distributed diazotroph Trichodesmium contributes importantly to nitrogen inputs in the oligotrophic oceans. Sites of dissolved organic matter (DOM) accumulation could promote the mixotrophic nutrition of Trichodesmium when inorganic nutrients are scarce. Nano-scale secondary ion mass spectrometry (nanoSIMS) analyses of individual trichomes sampled in the South Pacific Ocean, showed significant 13 C-enrichments after incubation with either 13 C-labeled carbohydrates or amino acids. These results suggest that DOM could be directly taken up by Trichodesmium or primarily consumed by heterotrophic epibiont bacteria that ultimately transfer reduced DOM compounds to their host trichomes. Although the addition of carbohydrates or amino acids did not significantly affect bulk N 2 fixation rates, N 2 fixation was enhanced by amino acids in individual colonies of Trichodesmium. We discuss the ecological advantages of DOM use by Trichodesmium as an alternative to autotrophic nutrition in oligotrophic open ocean waters. Nitrogen is recognized as the proximate limiting nutrient for primary production in the oceans 1. The oceanic nitrogen reservoir is controlled by a balance between fixed nitrogen gains (via dinitrogen-N 2-fixation) and losses (denitrification) 2. While global nitrogen budget estimations determine that denitrification exceeds N 2 fixation considerably 3 , recent improvements in the 15 N 2 isotope tracer method used to measure biological N 2 fixation have evidenced that formerly published rates could be underestimated by a factor of ~2 to 6 4–8 , and thus could be high enough to balance denitrification on a global basis. However, the variability among N 2 fixation rates obtained when using the two different methods (adding 15 N 2 as a bubble or pre-dissolved in seawater) 4,9 can be high 7 and at times not significant 10–12. A mechanistic understanding of which factors determine the degree of discrepancy between the two 15 N 2 methods is currently lacking. Moreover, marine pelagic N 2 fixation had been long attributed to the tropical and subtropical latitudinal bands of the ocean, e.g. 13 , while other ecological niches such as high latitude waters, oxygen minimum zones and the vast dark realm of the ocean (below the euphotic zone) are now recognized as active N 2 fixation sites 14–16. It is likely that the inclusion of these previously unaccounted for active N 2 fixation sites will be more important in equilibrating denitrification and N 2 fixation rates than the underestimation of rates due to discrepancies between isotopic tracer methods. In chronically stratified open ocean regions such as the vast subtropical gyres, primary production depends largely on external fixed nitrogen inputs provided by N 2 fixation performed by prokaryotes termed 'diazotrophs'. Diazotrophic cyanobacteria are photosynthetic prokaryotes (with the exception of the photoheterotrophic Candidatus Atelocyanobacterium thalassa which cannot photosynthesize) 17 that thrive in oligotrophic tropical and subtropical waters of the oceans where they provide an important source of fixed nitrogen for other phyto-plankton 13. Despite being classically regarded as photoautotrophs, some unicellular diazotrophic cyanobacteria like Cyanothece are able to take up dissolved organic matter (DOM) molecules photoheterotrophically 18. As well, various filamentous diazotrophic cyanobacteria such as Anabaena bear genes for amino acids transport, which may be used to incorporate amino acids from the in situ DOM pool, or to assimilate amino acids self-produced during diazotrophic growth 19

Distribution of normalized water-leaving radiances at UV and visible wave bands in relation with chlorophyll a and colored detrital matter content in the southeast Pacific

International audience[1] In-water radiometric measurements were performed in the southeast Pacific (8°S-35°S, 141°W-72°W) from October to December 2004 during the Biogeochemistry and Optics South Pacific Experiment cruise. Normalized water-leaving radiances (nL w (l)) were determined at eight wave bands within the ultraviolet (UV) (305, 325, 340, and 380 nm) and visible (412, 443, 490, and 565 nm) spectral domains. The highest nL w (l) (mW cm À2 sr À1) were recorded in the hyperoligotrophic waters of the South Pacific Gyre, with values increasing with wavelength from 305 (nL w = 0.64) to 380 nm (nL w = 3.18) in the UV range and decreasing from 412 (nL w = 4.46) to 565 nm (nL w = 0.23) in the visible region. The intense nL w (l) observed in the violet-blue domains were attributed to very low absorptions of colored detrital matter (CDM), likely related to a strong photobleaching of colored dissolved organic matter in the surface waters. We evaluated the relationships between the UV, violet, or blue/green wave band ratios of nL w (l) and surface total chlorophyll a (TChl a) concentration and CDM absorption (a CDM (l)). For TChl a, the best correlation was found with the blue/green ratio at 443 nm: TChl a (mg m À3) = 2.37[nL w (443)/nL w (565)] À1.51 (r 2 = 0.86 and RMS error (RMSE) = 23%). By contrast, for a CDM (l), the best correlation was observed when using the UV/green ratio at 325 nm: a CDM (325) (m À1) = 0.16[nL w (325)/nL w (565)] À0.69 (r 2 = 0.82 and RMSE = 16%). These results show the potential role of nL w (l) at UV wave bands for the assessment, through empirical algorithms, of colored detrital matter in the surface oceanic waters. Citation: Tedetti, M., B. Charrière, A. Bricaud, J. Para, P. Raimbault, and R. Sempéré (2010), Distribution of normalized water-leaving radiances at UV and visible wave bands in relation with chlorophyll a and colored detrital matter content in the southeast Pacific

Ecology and biogeochemistry of contrasting trophic environments in the South East Pacific by carbon isotope ratios on lipid biomarkers

International audienceThe distribution of lipid biomarkers and their carbon isotope composition was investigated on suspended particles from different contrasting trophic environments at six sites in the South East Pacific. High algal biomass with diatom-related lipids was characteristic in the upwelling zone, whereas haptophyte lipids were proportionally most abundant in the nutrient-poor settings of the centre of the South Pacific Gyre and on its easter edge. Dinoflagellate–sterols were minor contributors in all of the studied area and cyanobacteria-hydrocarbons were at maximum in the high nutrient low chlorophyll regime of the subequatorial waters at near the Marquesas archipelago. The taxonomic and spatial variability of the relationships between carbon photosynthetic fractionation and environmental conditions for four specific algal taxa (diatoms, haptophytes, dinoflagellates and cyanobacteria) was also investigated. The carbon isotope fractionation factor (ep) of the diatom marker varied over a range of 16‰ along the different trophic systems. In contrast, ep of dinoflagellate, cyanobacteria and alkenone markers varied only by 7–10‰. The low fractionation factors and small variations between the different phytoplankton markers measured in the upwelling area likely reveals uniformly high specific growth rates within the four phytoplankton taxa, and/or that transport of inorganic carbon into phytoplankton cells may not only occur by diffusion but by other carbon concentrating mechanisms (CCM). In contrast, in the oligotrophic zone, i.e. gyre and eastgyre, relatively high ep values, especially for the diatom marker, indicate diffusive CO2 uptake by the eukaryotic phytoplankton. At these nutrient-poor sites, the lowest ep values for haptophytes, dinoflagellates and cyanobacteria infer higher growth rates compared to diatoms

Nutrient dynamics and primary production in the eutrophic Berre Lagoon (Mediterranean, France)

1 - The brackish Berre Lagoon is one of the largest Mediterranean coastal lagoons (155 km2). Since 1966, it has been influenced by a large freshwater discharge from a hydroelectric power plant, which has led to strong changes in the ecosystem structure and functioning. 2 - During 2005 and 2006, we monitored physico-chemical variables, primary production, nitrogen assimilation and regeneration in the water column using the dual-isotopic 15N/13C technique. 3 - Significant seasonal variations were observed for salinity, however summer values were similar to those observed in the lagoon before the power plant opened (30). Nitrate concentrations (NO3-) varied, yielding very low values in the summer to values of 220 µg N-NO3-.1-1 in winter. Ammonium (NH4-) and phosphate (PO43-) were observed in significant concentrations throughout the year (>7 µg N-NH4+.l-1 and 3 µg P-PO43-.1-1 respectively). Total chlorophyll increased slightly from February 2005 to June 2006, never exceeding 17 µg.1-1 until summer 2006, which was marked by a large accumulation of biomass (> 30 µg.1-1) maintained until the end of the year. 4 - Mean primary production rates ranged from 50 to 1 600 µgC.1-1.d-1, giving a total annual production of 507 gC.m-2.yr-1 in 2005 and 742 gC.m-2.yr-1 in 2006. This production is based on the assimilation of 15 000 and 24 000 tons of dissolved inorganic nitrogen (DIN) in 2005 and 2006 respectively (uptake rates ranging from 3 to 77 µg N-NH4-+.1-1.d-1 and from 1 to 63 µg N-NO3-.1-1.d-1). Regeneration processes were very active: 15 to 85 µg N-NH4+.1-1.d-1 were regenerated in the water column and sustained at least 60% of ammonium uptake. 5 - 95% of the measured annual primary production was regenerated production. These budgets need to 5 - be analysed with caution as biological processes are characterized by high interannual and seasonal variabilities. Regeneration processes in the water column certainly fed by other processes (acting in the sediments) are the basis of eutrophication maintenance in the Berre Lagoon

Phosphate availability and the ultimate control of new nitrogen input by nitrogen fixation in the tropical Pacific Ocean

International audienceDue to the low atmospheric input of phosphate into the open ocean, it is one of the key nutrients that could ultimately control primary production and carbon export into the deep ocean. The observed trend over the last 20 years has shown a decrease in the dissolved inorganic phosphate (DIP) pool in the North Pacific gyre, which has been correlated to the increase in di-nitrogen (N2) fixation rates. Following a NW-SE transect, in the Southeast Pacific during the early austral summer (BIOSOPE cruise), we present data on DIP, dissolved organic phosphate (DOP) and particulate phosphate (PP) pools along with DIP turnover times (TDIP) and N2 fixation rates. We observed a decrease in DIP concentration from the edges to the centre of the gyre. Nevertheless the DIP concentrations remained above 100 nmol L-1 and T DIP was more than 6 months in the centre of the gyre; DIP availability remained largely above the level required for phosphate limitation to occur and the absence of Trichodesmium spp and low nitrogen fixation rates were likely to be controlled by other factors such as temperature or iron availability. This contrasts with recent observations in the North Pacific Ocean at the ALOHA station and in the western Pacific Ocean at the same latitude (DIAPALIS cruises) where lower DIP concentrations (-1) and T DIP 2 fixation rates and possibly carbon dioxide sequestration, if the primary ecophysiological controls, temperature and/or iron availability, were alleviated

Nutrients limitation of primary productivity in the Southeast Pacific (BIOSOPE cruise)

Revue sans Comité de lectureInternational audienceIron is an essential nutrient involved in a variety of biological processes in the ocean, including photosynthesis, respiration and nitrogen fixation. Atmospheric deposition of aerosols is recognized as the main source of iron for the surface ocean. In high nutrient, low chlorophyll areas, it is now clearly established that iron limits phytoplankton productivity but its biogeochemical role in low nutrient, low chlorophyll environments has been poorly studied. We investigated this question in the unexplored southeast Pacific, arguably the most oligotrophic area of the global ocean. Situated far from any continental aerosol source, the atmospheric iron flux to this province is amongst the lowest of the world ocean. Here we report that, despite low dissolved iron concentrations (~0.1 nmol l-1) measured across the whole gyre (3 stations situated in the center, the western and the eastern edge), photosynthesis and primary productivity are only limited by iron availability at the border of the gyre, but not in the center. The seasonal stability of the gyre has apparently allowed for the development of populations acclimated to these extreme oligotrophic conditions. Moreover, despite clear evidence of nitrogen limitation in the central gyre, we were unable to measure nitrogen fixation in our experiments, even after iron and/or phosphate additions, and cyanobacterial nifH gene abundances were extremely low compared to the North Pacific Gyre. The South Pacific gyre is therefore unique with respect to the physiological status of its phytoplankton populations