The composition and distribution of semi-labile dissolved organic matter across the southwest Pacific

The distribution and dynamics of dissolved organic carbon (DOC) and dissolved combined neutral sugars (DCNS) were studied across an increasing oligotrophic gradient (18 to 22∘&thinsp;S latitude) in the tropical South Pacific Ocean, spanning from the Melanesian Archipelago (MA) area to the western part of the South Pacific gyre (WGY), in austral summer as a part of the OUTPACE project. Our results show that DOC and DCNS concentrations exhibited no statistical differences between the MA and WGY areas (0–200&thinsp;m: 47–81&thinsp;µM&thinsp;C for DOC and 0.2-4.2&thinsp;µM&thinsp;C for DCNS). However, due to a deepening of the euphotic zone, a deeper penetration of DOC was noticeable at 150&thinsp;m of depth at the WGY area. Excess DOC (DOCEX) was determined as the difference between surface and deep-sea DOC values, and euphotic zone integrated stocks of both DOC and DOCEX were higher in the WGY than the MA area. Considering DOCEX as representative of semi-labile DOC (DOCSL), its residence time was calculated as the ratio of DOCSL to bacterial carbon demand (BCD). This residence time was 176±43 days (n=3) in the WGY area, about 3 times longer than in the MA area (Tr=51±13 days, n=8), suggesting an accumulation of semi-labile dissolved organic matter (DOM) in the surface waters of WGY. Average epipelagic (0–200&thinsp;m) DCNS yields (DCNS&thinsp;×&thinsp;DOC−1) based on volumetric data were roughly similar in both areas, accounting for ∼2.8&thinsp;% of DOC. DCNS exhibited a longer residence time in WGY (Tr=91±41 days, n=3) than in MA (Tr=31±10 days, n=8), further suggesting that this DCNS pool persists longer in the surface waters of the WGY. The accumulation of DOCEX in the surface waters of WGY is probably due to very slow bacterial degradation due to nutrient and/or energy limitation of heterotrophic prokaryotes, indicating that biologically produced DOC can be stored in the euphotic layer of the South Pacific gyre for a long period.</p

Dynamics and controls of heterotrophic prokaryotic production in the western tropical South Pacific Ocean: links with diazotrophic and photosynthetic activity

Heterotrophic prokaryotic production (BP) was studied in the western tropical South Pacific (WTSP) using the leucine technique, revealing spatial and temporal variability within the region. Integrated over the euphotic zone, BP ranged from 58 to 120 mg C m−2 d−1 within the Melanesian Archipelago, and from 31 to 50 mg C m−2 d−1 within the western subtropical gyre. The collapse of a bloom was followed during 6 days in the south of Vanuatu using a Lagrangian sampling strategy. During this period, rapid evolution was observed in the three main parameters influencing the metabolic state: BP, primary production (PP) and bacterial growth efficiency. With N2 fixation being one of the most important fluxes fueling new production, we explored relationships between BP, PP and N2 fixation rates over the WTSP. The contribution of N2 fixation rates to bacterial nitrogen demand ranged from 3 to 81 %. BP variability was better explained by the variability of N2 fixation rates than by that of PP in surface waters of the Melanesian Archipelago, which were characterized by N-depleted layers and low DIP turnover times (TDIP  100 h), deeper in the Melanesian Archipelago, or within the entire euphotic zone in the subtropical gyre. The bacterial carbon demand to gross primary production ratio ranged from 0.75 to 3.1. These values are discussed in the framework of various assumptions and conversion factors used to estimate this ratio, including the methodological errors, the daily variability of BP, the bacterial growth efficiency and one bias so far not considered: the ability for Prochlorococcus to assimilate leucine in the dark

Role of environmental factors for the vertical distribution (0–1000 m) of marine bacterial communities in the NW Mediterranean Sea

Bacterioplankton plays a central role in energy and matter fluxes in the sea, yet the factors that constrain its variation in marine systems are still poorly understood. Here we use the explanatory power of direct multivariate gradient analysis to evaluate the driving forces exerted by environmental parameters on bacterial community distribution in the water column. We gathered and analysed data from a one month sampling period from the surface to 1000 m depth at the JGOFS-DYFAMED station (NW Mediterranean Sea). This station is characterized by very poor horizontal advection currents which makes it an ideal model to test hypotheses on the causes of vertical stratification of bacterial communities. Capillary electrophoresis single strand conformation polymorphism (CE-SSCP) fingerprinting profiles analyzed using multivariate statistical methods demonstrated a vertical zonation of bacterial assemblages in three layers, above, in or just below the chlorophyll maximum and deeper, that remained stable during the entire sampling period. Through the use of direct gradient multivariate ordination analyses we demonstrate that a complex array of biogeochemical parameters is the driving force behind bacterial community structure shifts in the water column. Physico-chemical parameters such as phosphate, nitrate, salinity and to a lesser extent temperature, oxygen, dissolved organic carbon and photosynthetically active radiation acted in synergy to explain bacterial assemblages changes with depth. Analysis of lipid biomarkers of organic matter sources and fates suggested that bacterial community structure in the surface layers was in part explained by lipids of chloroplast origin. Further detailed analysis of pigment-based phytoplankton diversity gave evidence of a compartmentalized influence of several phytoplankton groups on bacterial community structure in the first 150 m depth

Inferring phytoplankton carbon and eco-physiological rates from diel cycles of spectral particulate beam-attenuation coefficient

The diurnal fluctuations in solar irradiance impose a fundamental frequency on ocean biogeochemistry. Observations of the ocean carbon cycle at these frequencies are rare, but could be considerably expanded by measuring and interpreting the inherent optical properties. A method is presented to analyze diel cycles in particulate beam-attenuation coefficient (&lt;i&gt;c&lt;/i&gt;&lt;sub&gt;p&lt;/sub&gt;) measured at multiple wavelengths. The method is based on fitting observations with a size-structured population model coupled to an optical model to infer the particle size distribution and physiologically relevant parameters of the cells responsible for the measured diel cycle in &lt;i&gt;c&lt;/i&gt;&lt;sub&gt;p&lt;/sub&gt;. Results show that the information related to size and contained in the spectral data can be exploited to independently estimate growth and loss rates during the day and night. In addition, the model can characterize the population of particles affecting the diel variability in &lt;i&gt;c&lt;/i&gt;&lt;sub&gt;p&lt;/sub&gt;. Application of this method to spectral &lt;i&gt;c&lt;/i&gt;&lt;sub&gt;p&lt;/sub&gt; measured at a station in the oligotrophic Mediterranean Sea suggests that most of the observed variations in &lt;i&gt;c&lt;/i&gt;&lt;sub&gt;p&lt;/sub&gt; can be ascribed to a synchronized population of cells with an equivalent spherical diameter around 4.6±1.5 μm. The inferred carbon biomass of these cells was about 5.2–6.0 mg m&lt;sup&gt;−3&lt;/sup&gt; and accounted for approximately 10% of the total particulate organic carbon. If successfully validated, this method may improve our in situ estimates of primary productivity

Deep silicon maxima in the stratified oligotrophic Mediterranean Sea

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-&lt;i&gt;a&lt;/i&gt; (TChl-&lt;i&gt;a&lt;/i&gt;) 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-&lt;i&gt;a&lt;/i&gt; (DCM) were evidenced during both seasons with maximal concentrations of 0.45 μmol L&lt;sup&gt;−1&lt;/sup&gt; for BSi, 0.26 μg L&lt;sup&gt;−1&lt;/sup&gt; for Fuco, and 1.70 μg L&lt;sup&gt;−1&lt;/sup&gt; for TChl-&lt;i&gt;a&lt;/i&gt;, 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 &lt;i&gt;Chaetoceros&lt;/i&gt; spp., &lt;i&gt;Leptocylindrus&lt;/i&gt; spp., &lt;i&gt;Pseudonitzschia&lt;/i&gt; spp. and the association between large centric diatoms (&lt;i&gt;Hemiaulus hauckii&lt;/i&gt; and &lt;i&gt;Rhizosolenia styliformis&lt;/i&gt;) and the cyanobacterium &lt;i&gt;Richelia intracellularis&lt;/i&gt; 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

Inferring phytoplankton carbon and eco-physiological rates from diel cycles of spectral particulate beam-attenuation coefficient

The diurnal fluctuations in solar irradiance impose a fundamental frequency on ocean biogeochemistry. Observations of the ocean carbon cycle at these frequencies are rare, but could be considerably expanded by measuring and interpreting the inherent optical properties. A method is presented to analyze diel cycles in particulate beam-attenuation coefficient (cp) measured at multiple wavelengths. The method is based on fitting observations with a size-structured population model coupled to an optical model to infer the particle size distribution and physiologically relevant parameters of the cells responsible for the measured diel cycle in cp. Results show that the information related to size and contained in the spectral data can be exploited to independently estimate growth and loss rates during the day and night. In addition, the model can characterize the population of particles affecting the diel variability in cp. Application of this method to spectral cp measured at a station in the oligotrophic Mediterranean Sea suggests that most of the observed variations in cp can be ascribed to a synchronized population of cells with an equivalent spherical diameter around 4.6-1.5 1/4μm. The inferred carbon biomass of these cells was about 5.2-6.0 mg mg -\u273 and accounted for approximately 10% of the total particulate organic carbon. If successfully validated, this method may improve our in situ estimates of primary productivity

Inferring phytoplankton carbon and eco-physiological rates from diel cycles of spectral particulate beam-attenuation coefficient

The diurnal fluctuations in solar irradiance impose a fundamental frequency on ocean biogeochemistry. Observations of the ocean carbon cycle at these frequencies are rare, but could be considerably expanded by measuring and interpreting the inherent optical properties. A method is presented to analyze diel cycles in particulate beam-attenuation coefficient (c[subscript p]) measured at multiple wavelengths. The method is based on fitting observations with a size-structured population model coupled to an optical model to infer the particle size distribution and physiologically relevant parameters of the cells responsible for the measured diel cycle in c[subscript p]. Results show that the information related to size and contained in the spectral data can be exploited to independently estimate growth and loss rates during the day and night. In addition, the model can characterize the population of particles affecting the diel variability in c[subscript p]. Application of this method to spectral c[subscript p] measured at a station in the oligotrophic Mediterranean Sea suggests that most of the observed variations in c[subscript p] can be ascribed to a synchronized population of cells with an equivalent spherical diameter around 4.6±1.5 μm. The inferred carbon biomass of these cells was about 5.2–6.0 mg m⁻³ and accounted for approximately 10% of the total particulate organic carbon. If successfully validated, this method may improve our in situ estimates of primary productivity

Seasonal and interannual variability of the pelagic ecosystem and of the organic carbon budget in the Rhodes Gyre (eastern Mediterranean): influence of winter mixing

The Rhodes Gyre is a cyclonic persistent feature of the general circulation of the Levantine Basin in the eastern Mediterranean Sea. Although it is located in the most oligotrophic basin of the Mediterranean Sea, it is a relatively high primary production area due to strong winter nutrient supply associated with the formation of Levantine Intermediate Water. In this study, a 3D coupled hydrodynamic–biogeochemical model (SYMPHONIE/Eco3M-S) was used to characterize the seasonal and interannual variability of the Rhodes Gyre's ecosystem and to estimate an annual organic carbon budget over the 2013–2020 period. Comparisons of model outputs with satellite data and compiled in situ data from cruises and Biogeochemical-Argo floats revealed the ability of the model to reconstruct the main seasonal and spatial biogeochemical dynamics of the Levantine Basin. The model results indicated that during the winter mixing period, phytoplankton first progressively grow sustained by nutrient supply. Then, short episodes of convection driven by heat loss and wind events, favoring nutrient injections, organic carbon export, and inducing light limitation on primary production, alternate with short episodes of phytoplankton growth. The estimate of the annual organic carbon budget indicated that the Rhodes Gyre is an autotrophic area, with a positive net community production in the upper layer (0–150 m) amounting to 31.2 ± 6.9 gCm-2yr-1. Net community production in the upper layer is almost balanced over the 7-year period by physical transfers, (1) via downward export (16.8 ± 6.2 gCm-2yr-1) and (2) through lateral transport towards the surrounding regions (14.1 ± 2.1 gCm-2yr-1). The intermediate layer (150–400 m) also appears to be a source of organic carbon for the surrounding Levantine Sea (7.5 ± 2.8 gCm-2yr-1) mostly through the subduction of Levantine Intermediate Water following winter mixing. The Rhodes Gyre shows high interannual variability with enhanced primary production, net community production, and exports during years marked by intense heat losses and deep mixed layers. However, annual primary production appears to be only partially driven by winter vertical mixing. Based on our results, we can speculate that future increase of temperature and stratification could strongly impact the carbon fluxes in this region.</p

Taxonomic and Environmental Variability in the Elemental Composition and Stoichiometry of Individual Dinoflagellate and Diatom Cells from the NW Mediterranean Sea

Here we present, for the first time, the elemental concentration, including C, N and O, of single phytoplankton cells collected from the sea. Plankton elemental concentration and stoichiometry are key variables in phytoplankton ecophysiology and ocean biogeochemistry, and are used to link cells and ecosystems. However, most field studies rely on bulk techniques that overestimate carbon and nitrogen because the samples include organic matter other than plankton organisms. Here we used X-ray microanalysis (XRMA), a technique that, unlike bulk analyses, gives simultaneous quotas of C, N, O, Mg, Si, P, and S, in single-cell organisms that can be collected directly from the sea. We analysed the elemental composition of dinoflagellates and diatoms (largely Chaetoceros spp.) collected from different sites of the Catalan coast (NW Mediterranean Sea). As expected, a lower C content is found in our cells compared to historical values of cultured cells. Our results indicate that, except for Si and O in diatoms, the mass of all elements is not a constant fraction of cell volume but rather decreases with increasing cell volume. Also, diatoms are significantly less dense in all the measured elements, except Si, compared to dinoflagellates. The N:P ratio of both groups is higher than the Redfield ratio, as it is the N:P nutrient ratio in deep NW Mediterranean Sea waters (N:P = 20–23). The results suggest that the P requirement is highest for bacterioplankton, followed by dinoflagellates, and lowest for diatoms, giving them a clear ecological advantage in P-limited environments like the Mediterranean Sea. Finally, the P concentration of cells of the same genera but growing under different nutrient conditions was the same, suggesting that the P quota of these cells is at a critical level. Our results indicate that XRMA is an accurate technique to determine single cell elemental quotas and derived conversion factors used to understand and model ocean biogeochemical cycles

Variabilité hydrologique et biologique du golfe du Lion. I. Transports en azote et productivité potentielle

A monthly time-series of the seasonal evolution of the North Mediterranean Current (NMC) and of nitrogen flux across a transect (5 degrees 12'5 E between 43 degrees 10'N and 42 degrees 50'N) at the entrance of the Gulf of Lions was carried out over two years. The NMC exhibited two distinct structures: a winter structure (from November to May) characterised by strong mesoscale and interannual variations in which the current was 20 to 30 km width, about 400 m depth and velocity scaled between 5 and more than 40 cm s(-1); and a summer structure (from May to November) which was less sensitive to mesoscale variations but showing large interannual variability. The current was located more offshore, was wider and shallower than the one in winter and velocity remained below 20 cm s(-1). Meteorological conditions, frontal structures on both sides of the NMC, coastal upwelling and continental inputs explained the observed mesoscale discrepancies between water flux and nitrogen transport. Nitrogen input from the general circulation across the prospected transect is estimated at 187 +/- 20 kT a(-1) in the 0-200 m layer, with a distribution of 39% nitrate, 6% particulate nitrogen and 55% dissolved organic nitrogen. Nitrate input (25 +/- 3 kT a(-1)) in the 0-100 m upper layer could sustain more than one third of the potential new production in the Gulf of Lions.Un suivi mensuel de l'évolution saisonnière du Courant Nord Méditerranéen (CNM) et des flux en azote à travers une radiale (5° 12′5 E entre 43° 10′N et 42° 50′N) a été réalisé, à l'entrée du golfe du Lion, sur une période de deux ans. Le CNM est décrit selon deux structures distinctes. Une structure hivernale (de novembre à mai) caractérisée par de fortes variations à méso-échelle et interannuelles; la veine de courant est large de 20 à 30 km et profonde d'environ 400 m, avec des vitesses comprises entre 5 et plus de 40 cm s−1. Une structure estivale (entre mai et novembre) moins sensible aux variations à méso-échelle mais présentant une forte variabilité interannuelle; la veine est alors plus hauturière, plus large et moins profonde qu'en hiver, les vitesses restant généralement inférieures à 20 cm s−1. Les conditions météorologiques, la présence de zones frontales sur les bords interne et externe du CNM, les phénomènes d'upwelling côder et d'apports continentaux permettent d'expliquer les découplages observés entre les flux en eau et en azote à travers la radiale. Les apports en azote par la circulation générale à travers la radiale sont estimés à 187 ± 20 kT a−1 dans la couche 0–200 m, et se répartissent en 39 % de nitrate, 6 % d'azote particulaire et 55 % d'azote organique dissous. Les apports de nitrate dans la couche superficielle 0–100 m (25 ± 3 kTa−1) pourraient être à l'origine de près d'un tiers de la production nouvelle potentielle du golfe du Lion