Search

Hydrolysis of dissolved organic phosphorus by marine planktonic microorganisms is a key process in the P cycle, particularly in P-depleted, oligotrophic environments. The present study assessed spatiotemporal variations in phosphomonoesterase (PME) and phosphodiesterase (PDE) activities using concentration kinetics in the eastern Mediterranean Sea in two contrasting situations: the end of winter (including a small bloom period) and autumn. The distribution and regulation of the maximum hydrolysis rate (Vm) and half-saturation constant (Km) of both ectoenzymes were assessed in relation to the vertical structure of the epipelagic layers. PME reached its maximum activities (Vm) after the addition of 1 µM MUF-P (4-methylumbelliferyl phosphate), whereas, for PDE, it was necessary to add up to 50 µM bis(4-methylumbelliferyl)phosphate (bis-MUF-P) to reach saturation state. On average, the Km of PDE was 33 ± 25 times higher than that of PME. The Vm of PME and Vm of PDE were linearly correlated. Conversely to the Km values, Vm values were on the same order of magnitude for both ectoenzymes, with their ratio (Vm PME : Vm PDE) ranging between 0.2 and 6.3. Dissolved organic phosphorus (DOP) and the phosphomonoesterase hydrolysable fraction of DOP explained most of the lack of variability in Vm PME and Vm PDE. On the contrary, Vm of both phosphohydrolase enzymes was inversely correlated to the concentration of dissolved inorganic phosphorus. The particular characteristics of concentration kinetics obtained for PDE (saturation at 50 µM, high Km, high turnover times) are discussed with respect to the possible unequal distribution of PDE and PME among the size continuum of organic material and accessibility of phosphodiesters.

Total alkalinity (AT) and dissolved inorganic carbon (CT) in the oceans are important properties with respect to understanding the ocean carbon cycle and its link to global change (ocean carbon sinks and sources, ocean acidification) and ultimately finding carbon-based solutions or mitigation procedures (marine carbon removal). We present a database of more than 44 400 AT and CT observations along with basic ancillary data (spatiotemporal location, depth, temperature and salinity) from various ocean regions obtained, mainly in the framework of French projects, since 1993. This includes both surface and water column data acquired in the open ocean, coastal zones and in the Mediterranean Sea and either from time series or dedicated one-off cruises. Most AT and CT data in this synthesis were measured from discrete samples using the same closed-cell potentiometric titration calibrated with Certified Reference Material, with an overall accuracy of ±4 µmol kg−1 for both AT and CT. The data are provided in two separate datasets – for the Global Ocean and the Mediterranean Sea (https://doi.org/10.17882/95414, Metzl et al., 2023), respectively – that offer a direct use for regional or global purposes, e.g., AT–salinity relationships, long-term CT estimates, and constraint and validation of diagnostic CT and AT reconstructed fields or ocean carbon and coupled climate–carbon models simulations as well as data derived from Biogeochemical-Argo (BGC-Argo) floats. When associated with other properties, these data can also be used to calculate pH, the fugacity of CO2 (fCO2) and other carbon system properties to derive ocean acidification rates or air–sea CO2 fluxes.

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.

According to current experimental evidence, coral reefs could disappear within the century if CO2 emissions remain unabated. However, recent discoveries of diverse and high cover reefs that already live under extreme conditions suggest that some corals might thrive well under hot, high-pCO2, and deoxygenated seawater. Volcanic CO2 vents, semi-enclosed lagoons, and mangrove estuaries are unique study sites where one or more ecologically relevant parameters for life in the oceans are close to or even worse than currently projected for the year 2100. Although they do not perfectly mimic future conditions, these natural laboratories offer unique opportunities to explore the mechanisms that reef species could use to keep pace with climate change. To achieve this, it is essential to characterize their environment as a whole and accurately consider all possible environmental factors that may differ from what is expected in the future, possibly altering the ecosystem response. This study focuses on the semi-enclosed lagoon of Bouraké (New Caledonia, southwest Pacific Ocean) where a healthy reef ecosystem thrives in warm, acidified, and deoxygenated water. We used a multi-scale approach to characterize the main physical-chemical parameters and mapped the benthic community composition (i.e., corals, sponges, and macroalgae). The data revealed that most physical and chemical parameters are regulated by the tide, strongly fluctuate three to four times a day, and are entirely predictable. The seawater pH and dissolved oxygen decrease during falling tide and reach extreme low values at low tide (7.2 pHT and 1.9 mg O2 L−1 at Bouraké vs. 7.9 pHT and 5.5 mg O2 L−1 at reference reefs). Dissolved oxygen, temperature, and pH fluctuate according to the tide by up to 4.91 mg O2 L−1, 6.50 ∘C, and 0.69 pHT units on a single day. Furthermore, the concentration of most of the chemical parameters was 1 to 5 times higher at the Bouraké lagoon, particularly for organic and inorganic carbon and nitrogen but also for some nutrients, notably silicates. Surprisingly, despite extreme environmental conditions and altered seawater chemical composition measured at Bouraké, our results reveal a diverse and high cover community of macroalgae, sponges, and corals accounting for 28, 11, and 66 species, respectively. Both environmental variability and nutrient imbalance might contribute to their survival under such extreme environmental conditions. We describe the natural dynamics of the Bouraké ecosystem and its relevance as a natural laboratory to investigate the benthic organism's adaptive responses to multiple extreme environmental conditions.

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∘ 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 m: 47–81 µM C for DOC and 0.2-4.2 µM C for DCNS). However, due to a deepening of the euphotic zone, a deeper penetration of DOC was noticeable at 150 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 m) DCNS yields (DCNS × DOC−1) based on volumetric data were roughly similar in both areas, accounting for ∼2.8 % 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.

The fate of diazotroph (N2 fixers) derived carbon (C) and nitrogen (N) and their contribution to vertical export of C and N in the western tropical South Pacific Ocean was studied during OUTPACE (Oligotrophy to UlTra-oligotrophy PACific Experiment). Our specific objective during OUTPACE was to determine whether autocatalytic programmed cell death (PCD), occurring in some diazotrophs, is an important mechanism affecting diazotroph mortality and a factor regulating the vertical flux of organic matter and, thus, the fate of the blooms. We sampled at three long duration (LD) stations of 5 days each (LDA, LDB and LDC) where drifting sediment traps were deployed at 150, 325 and 500 m depth. LDA and LDB were characterized by high chlorophyll a (Chl a) concentrations (0.2–0.6 µg L−1) and dominated by dense biomass of the filamentous cyanobacterium Trichodesmium as well as UCYN-B and diatom–diazotroph associations (Rhizosolenia with Richelia-detected by microscopy and het-1 nifH copies). Station LDC was located at an ultra-oligotrophic area of the South Pacific gyre with extremely low Chl a concentration ( ∼ 0.02 µg L−1) with limited biomass of diazotrophs predominantly the unicellular UCYN-B. Our measurements of biomass from LDA and LDB yielded high activities of caspase-like and metacaspase proteases that are indicative of PCD in Trichodesmium and other phytoplankton. Metacaspase activity, reported here for the first time from oceanic populations, was highest at the surface of both LDA and LDB, where we also obtained high concentrations of transparent exopolymeric particles (TEP). TEP were negatively correlated with dissolved inorganic phosphorus and positively coupled to both the dissolved and particulate organic carbon pools. Our results reflect the increase in TEP production under nutrient stress and its role as a source of sticky carbon facilitating aggregation and rapid vertical sinking. Evidence for bloom decline was observed at both LDA and LDB. However, the physiological status and rates of decline of the blooms differed between the stations, influencing the amount of accumulated diazotrophic organic matter and mass flux observed in the traps during our experimental time frame. At LDA sediment traps contained the greatest export of particulate matter and significant numbers of both intact and decaying Trichodesmium, UCYN-B and het-1 compared to LDB where the bloom decline began only 2 days prior to leaving the station and to LDC where no evidence for bloom or bloom decline was seen. Substantiating previous findings from laboratory cultures linking PCD to carbon export in Trichodesmium, our results from OUTPACE indicate that nutrient limitation may induce PCD in high biomass blooms such as displayed by Trichodesmium or diatom–diazotroph associations. Furthermore, PCD combined with high TEP production will tend to facilitate cellular aggregation and bloom termination and will expedite vertical flux to depth.

The western tropical South Pacific (WTSP) Ocean has been recognized as a global hot spot of dinitrogen (N2) fixation. Here, as in other marine environments across the oceans, N2 fixation studies have focused on the sunlit layer. However, studies have confirmed the importance of aphotic N2 fixation activity, although until now only one had been performed in the WTSP. In order to increase our knowledge of aphotic N2 fixation in the WTSP, we measured N2 fixation rates and identified diazotrophic phylotypes in the mesopelagic layer along a transect spanning from New Caledonia to French Polynesia. Because non-cyanobacterial diazotrophs presumably need external dissolved organic matter (DOM) sources for their nutrition, we also identified DOM compounds using Fourier transform ion cyclotron resonance mass spectrometry (FTICRMS) with the aim of searching for relationships between the composition of DOM and non-cyanobacterial N2 fixation in the aphotic ocean. N2 fixation rates were low (average 0.63 ± 0.07 nmol N L−1 d−1) but consistently detected across all depths and stations, representing ∼ 6–88 % of photic N2 fixation. N2 fixation rates were not significantly correlated with DOM compounds. The analysis of nifH gene amplicons revealed a wide diversity of non-cyanobacterial diazotrophs, mostly matching clusters 1 and 3. Interestingly, a distinct phylotype from the major nifH subcluster 1G dominated at 650 dbar, coinciding with the oxygenated Subantarctic Mode Water (SAMW). This consistent pattern suggests that the distribution of aphotic diazotroph communities is to some extent controlled by water mass structure. While the data available are still too scarce to elucidate the distribution and controls of mesopelagic non-cyanobacterial diazotrophs in the WTSP, their prevalence in the mesopelagic layer and the consistent detection of active N2 fixation activity at all depths sampled during our study suggest that aphotic N2 fixation may contribute significantly to fixed nitrogen inputs in this area and/or areas downstream of water mass circulation.

Surface waters (0–200 m) of the western tropical South Pacific (WTSP) were sampled along a longitudinal 4000 km transect (OUTPACE cruise, DOI: 10.17600/15000900) during the austral summer (stratified) period (18 February to 3 April 2015) between the Melanesian Archipelago (MA) and the western part of the SP gyre (WGY). Two distinct areas were considered for the MA, the western MA (WMA), and the eastern MA (EMA). The main carbon (C), nitrogen (N), and phosphorus (P) pools and fluxes provide a basis for the characterization of the expected trend from oligotrophy to ultra-oligotrophy, and the building of first-order budgets at the daily and seasonal timescales (using climatology). Sea surface chlorophyll a well reflected the expected oligotrophic gradient with higher values obtained at WMA, lower values at WGY, and intermediate values at EMA. As expected, the euphotic zone depth, the deep chlorophyll maximum, and nitracline depth deepen from west to east. Nevertheless, phosphaclines and nitraclines did not match. The decoupling between phosphacline and nitracline depths in the MA allows for excess P to be locally provided in the upper water by winter mixing. We found a significant biological soft tissue carbon pump in the MA sustained almost exclusively by dinitrogen (N2) fixation and essentially controlled by phosphate availability in this iron-rich environment. The MA appears to be a net sink for atmospheric CO2, while the WGY is in quasi-steady state. We suggest that the necessary excess P, allowing the success of nitrogen fixers and subsequent carbon production and export, is mainly brought to the upper surface by local deep winter convection at an annual timescale rather than by surface circulation. While the origin of the decoupling between phosphacline and nitracline remains uncertain, the direct link between local P upper water enrichment, N2 fixation, and organic carbon production and export, offers a possible shorter timescale than previously thought between N input by N2 fixation and carbon export. The low iron availability in the SP gyre and P availability in the MA during the stratified period may appear as the ultimate control of N input by N2 fixation. Because of the huge volume of water to consider, and because the SP Ocean is the place of intense denitrification in the east (N sink) and N2 fixation in the west (N source), precise seasonal C, N, P, and iron (Fe) budgets would be of prime interest to understand the efficiency, at the present time and in the future, of the oceanic biological carbon pump.

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 2 fixation rates where DIP was more available (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.

The 2009–2010 period was marked by an episode of intense drought known as the El Niño Modoki event. Sampling of the Términos Lagoon (Mexico) was carried out in November 2009 in order to understand the influence of these particular environmental conditions on organic matter fluxes within the lagoon's pelagic ecosystem and, more specifically, on the relationship between phyto- and bacterioplankton communities. The measurements presented here concern biogeochemical parameters (nutrients, dissolved and particulate organic matter [POM], and dissolved polycyclic aromatic hydrocarbons [PAHs]), phytoplankton (biomass and photosynthesis), and bacteria (diversity and abundance, including PAH degradation bacteria and ectoenzymatic activities). During the studied period, the water column of the Términos Lagoon functioned globally as a sink and, more precisely, as a nitrogen assimilator. This was due to the high production of particulate and dissolved organic matter (DOM), even though exportation of autochthonous matter to the Gulf of Mexico was weak. We found that bottom-up control accounted for a large portion of the variability of phytoplankton productivity. Nitrogen and phosphorus stoichiometry mostly accounted for the heterogeneity in phytoplankton and free-living prokaryote distribution in the lagoon. In the eastern part, we found a clear decoupling between areas enriched in dissolved inorganic nitrogen near the Puerto Real coastal inlet and areas enriched in phosphate (PO4) near the Candelaria estuary. Such a decoupling limited the potential for primary production, resulting in an accumulation of dissolved organic carbon and nitrogen (DOC and DON, respectively) near the river mouths. In the western part of the lagoon, maximal phytoplankton development resulted from bacterial activity transforming particulate organic phosphorus (PP) and dissolved organic phosphorus (DOP) to available PO4 and the coupling between Palizada River inputs of nitrate (NO3) and PP. The Chumpan River contributed only marginally to PO4 inputs due to its very low contribution to overall river inputs. The highest dissolved total PAH concentrations were measured in the El Carmen Inlet, suggesting that the anthropogenic pollution of the zone is probably related to the oil-platform exploitation activities in the shallow waters of the southern of the Gulf of Mexico. We also found that a complex array of biogeochemical and phytoplanktonic parameters were the driving force behind the geographical distribution of bacterial community structure and activities. Finally, we showed that nutrients brought by the Palizada River supported an abundant bacterial community of PAH degraders, which are of significance in this important oil-production zone.

Intact polar membrane lipids compose a significant fraction of cellular material in plankton and their synthesis imposes a substantial constraint on planktonic nutrient requirements. As a part of the Biogeochemistry from the Oligotrophic to the Ultraoligotrophic Mediterranean (BOUM) cruise we examined the distribution of several classes of intact polar diacylglycerolipids (IP-DAGs) across the Mediterranean, and found that phospholipid concentration as a percent of total lipids correlated with phosphate concentration. In addition, the ratios of non-phosphorus lipids to phospholipids – sulfoquinovosyldiacylglycerol (SQDG) to phosphatidylglycerol (PG), and betaine lipids to phosphatidylcholine (PC) – were also found to increase from west to east across the Mediterranean. Additionally, microcosm incubations from across the Mediterranean were amended with phosphate and ammonium, and in the course of several days nutrient amendments elicited a shift in the ratios of IP-DAGs. These experiments were used to assess the relative contribution of community shifts and physiological response to the observed change in IP-DAGs across the Mediterranean. The ratio of SQDG to chlorophyll-a was also explored as an indicator of phytoplankton response to nitrogen availability. This study is the first to demonstrate the dynamic response of membrane lipid composition to changes in nutrients in a natural, mixed planktonic community.

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 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.

A coupled hydrodynamic-biogeochemical modelling is developed to address main mechanisms that drive the particulate organic carbon (POC) deposition in the Gulf of Lions (NW-Mediterranean). Low-salinity water (LSW, salinity

The variability of inherent optical properties is investigated in the ultra-oligotrophic waters of the Mediterranean Sea sampled during the BOUM experiment performed during early summer 2008. Bio-optical relationships found for ultra-oligotrophic waters of the three anticyclonic gyres sampled significantly depart from the mean standard relationships provided for the global ocean, confirming the peculiar character of these Mediterranean waters. These optical anomalies are diversely related to the specific biological and environmental conditions occurring in the studied ecosystem. Specifically, the surface specific phytoplankton absorption coefficient exhibits values lower than those expected from the general relationships mainly in relation with a high contribution of relatively large sized phytoplankton. Conversely, the particulate backscattering coefficient, bbp, values are much higher than the mean standard values for a given chlorophyll-a concentration, TChl-a. This feature can presumably be related to the relevant influence of highly refractive submicrometer particles of Saharan origin in the surface layer of the water column. The present measurements also show that the Mediterranean Sea is greener than TChl-a alone indicates, as already stressed in previous studies. This color anomaly is partly explained by the estimated colored dissolved organic matter and submicrometer particles absorption coefficients, and to a greater extent by the high bbp/TChl-a values assuming that these particles backscatter light similarly in the green and blue parts of the visible spectrum. The diel variation of both the particulate matter attenuation and backscattering coefficients were also investigated specifically. Despite some differences in the timing and the magnitude of the daily oscillations found for these optical parameters, potential for the backscattering coefficient daily oscillation to be used, similarly to that for the attenuation coefficient, as a proxy for estimating carbon community production budget has been highlighted for the first time. This result is particularly relevant for present and future geostationary spatial ocean color missions.

A better understanding of the factors controlling N2 fixation is a pre-requisite for improving our knowledge on the contribution of N2 fixation process in the nitrogen cycling. Trace-metal clean nutrient/dust addition bioassays (+P, +PFe, +dust) were performed at three stations located in the western, central and eastern Mediterranean Sea, in summer 2008 as part of the BOUM cruise. The main goals were (1) to investigate the nutrient factor(s) limiting N2 fixation (uptake of 15N2) and (2) to evaluate the potential impact of a Saharan dust event on this biological process during the stratification period. Initially, surface waters at the three stations were DIP-depleted (2 fixation (from 130 % to 430 %). The highest dust stimulation of N2 fixation was recorded at the station located in the eastern basin. The response of diazotrophic activity to nutrient additions was variable between the sampled stations suggesting a spatial variability of the factor controlling N2 fixation over the whole basin. At all stations, N2 fixation was not limited by Fe nor co-limited by P and Fe. At the western station, N2 fixation was DIP limited while at the eastern one, N2 fixation was first DIP limited, then was limited by one or several chemical element(s) released by dust. Our results demonstrated that a Saharan dust input was able to relieve these successive on going limitations. Very interestingly, at the station located in the central basin, N2 fixation was not limited by the availability of P yet it was strongly stimulated by dust addition (x3.1). A chemical element or a combination of several, released by the added dust may have been responsible for the observed stimulations of N2 fixation. These results indicated that Saharan dust pulses to the surface Mediterranean waters, in addition to P and Fe, could be a source of chemical(s) element(s) that are necessary for metabolic processes and therefore influence rates of N2 fixation.

We examined the effects of nutrient additions on rates of 14C-based particulate and dissolved primary production as well as O2-based metabolic rates in surface waters (8 m) of three anticyclonic eddies, located in the Western, Central and Eastern Mediterranean. Ship-board microcosm experiments employing additions of inorganic nitrogen (+N) and phosphorus (+P), alone and in combination (+NP), were conducted in June/July 2008 during the BOUM (Biogeochemistry from the Oligotrophic to the Ultra-oligotrophic Mediterranean) cruise. In all three experiments, particulate primary production was significantly stimulated by the additions of nitrogen (+N, +NP) while no effect was observed with the addition of phosphorus alone (+P). Percent extracellular release of photosynthate (PER) displayed the lowest values (4–8 %) in the +NP treatment. Among the three treatments (+N, +P, +NP), the +NP had the strongest effect on oxygen metabolic rates, leading to positive values of net community production (NCP > 0). These changes of NCP were mainly due to enhanced gross primary production (GPP) rather than reduced dark community respiration rates (DCR). In all three sites, in +NP treatment autotrophic production (whether expressed as GPP or PPtotal) was sufficient to fulfil the estimated carbon requirements of heterotrophic prokaryotes, while addition of nitrogen alone (+N) had a weaker effect on GPP, resulting in metabolically balanced systems. At the three sites, in treatments with N (+N, +NP), phytoplankton and heterotrophic prokaryote production were positively correlated. Heterotrophic conditions were observed in the Control and +P treatment at the central and eastern sites, and autotrophic production was not sufficient to supply estimated bacterial carbon demand, evidence of a decoupling of phytoplankton production and consumption by heterotrophic prokaryotes.

The abundance and activity of the major members of the heterotrophic microbial community – from viruses to ciliates – were studied along a longitudinal transect across the Mediterranean Sea in the summer of 2008. The Mediterranean Sea is characterized by a west to-east gradient of deepening of DCM (deep chlorophyll maximum) and increasing oligotrophy reflected in gradients of biomass and production. However, within this well documented longitudinal trend, hydrological mesoscale features exist and likely influence microbial dynamics. Here we present data from a W-E transect of 17 stations during the period of summer stratification. Along the transect the production and fate of organic matter was investigated at three selected sites each one located in the centre of an anticyclonic eddy: in the Algero-Provencal Basin (St. A), the Ionian Basin (St. B), and the Levantine Basin (St. C). The 3 geographically distant eddies showed low values of the different heterotrophic compartments of the microbial food web, and except for viruses in site C, all integrated (0–150 m) stocks were higher in reference stations located in the same basin outside the eddies. During our study the 3 eddies showed equilibrium between GPP (Gross Primary Production) and DCR (Dark Community Respiration). Integrated PPp (Particulate Primary Production) values at A, B and C varied from ~140 to ~190 mg C m−2.

Heterotrophic bacterioplankton abundance and production were investigated with depth (down to bathypelagic layers) and with longitude (from 4.9° E to 32.7° E) along a cruise track across the Mediterranean Sea in early summer 2008. Abundances and flow cytometric characteristics (green fluorescence and side scatter signals) of high nucleic acid (HNA) and low nucleic acid (LNA) bacterial cells were determined using flow cytometry. Contrary to what is generally observed, the relative importance of HNA cells, as a percent of total cells, (%HNA, range 30–69 %) was inversely related to bacterial production (range 0.15–44 ng C l−1 h−1) although the negative relation was weak (log–log regression r2=0.19). The %HNA as well as the mean side scatter of HNA group increased significantly with depth in the meso and bathypelagic layers. Vertical stratification played an important role in influencing the distribution and characteristics of bacterial cells especially with regard to layers located above, within or below the deep chlorophyll maximum. Within a given layer, the relationships between the flow cytometric characteristics and environmental variables such as chlorophyll-a, nutrients or bacterial production changed. Overall, the relationships between HNA and LNA cells and environmental parameters differed vertically more than longitudinally.

This paper provides an extensive vertical and longitudinal description of the biogeochemistry along an East-West transect of 3000 km across the Mediterranean Sea during summer 2008 (BOUM cruise). During this period of strong stratification, the distribution of nutrients, particulate and dissolved organic carbon (DOC), nitrogen (DON) and phosphorus (DOP) were examined to produce a detailed spatial and vertically extended description of the elemental stoichiometry of the Mediterranean Sea. Surface waters were depleted in nutrients and the thickness of this depleted layer increased towards the East from about 10 m in the Gulf of Lion to more than 100 m in the Levantine basin, with the phosphacline deepening to a greater extent than that for corresponding nitracline and thermocline depths. We used the minimum oxygen concentration through the water column in combination with 2 fixed concentrations of dissolved oxygen to distinguish an intermediate layer (Mineralization Layer; ML) from surface (Biogenic Layer; BL), and deep layers (DL). Whilst each layer was represented by different water masses, this approach allowed us to propose a schematic box-plot representation of the biogeochemical functioning of the two Mediterranean basins. Despite the increasing oligotrophic nature and the degree of P-depletion along the West to East gradient strong similarities were encountered between eastern and western ecosystems. Within the BL, the C:N:P ratios in all pools largely exceeded the Redfield ratios, but surprisingly, the nitrate vs. phosphate ratios in the ML and DL tended towards the canonical Redfield values in both basins. A change in particulate matter composition has been identified by a C increase relative to N and P along the whole water column in the western basin and between BL and ML in the eastern one. Our data showed a noticeable stability of the DOC:DON ratio (12–13) throughout the Mediterranean Sea. This is in good agreement with a P-limitation of microbial activities but in contradiction of the accepted concept that N is recycled faster than C. The western and eastern basins had similar or close biological functioning. Differences come from variability in the allochtonous nutrient sources in terms of quantity and quality, and to the specific hydrodynamic features of the Mediterranean basins.

Aerobic anoxygenic phototrophic (AAP) bacteria are photoheterotrophic prokaryotes able to use both light and organic substrates for energy production. They are widely distributed in coastal and oceanic environments and may contribute significantly to the carbon cycle in the upper ocean. To better understand questions regarding links between the ecology of these photoheterotrophic bacteria and the trophic status of water masses, we examined their horizontal and vertical distribution and the effects of nutrient additions on their growth along an oligotrophic gradient in the Mediterranean Sea. Concentrations of bacteriochlorophyll-a (BChl-a) and AAP bacterial abundance decreased from the western to the eastern basin of the Mediterranean Sea and were linked with concentrations of chlorophyll-a, nutrient and dissolved organic carbon. Inorganic nutrient and glucose additions to surface seawater samples along the oligotrophic gradient revealed that AAP bacteria were nitrogen- and carbon-limited in the ultraoligotrophic eastern basin. The intensity of the AAP bacterial growth response generally differed from that of the total bacterial growth response. BChl-a quota of AAP bacterial communities was significantly higher in the eastern basin than in the western basin, suggesting that reliance on phototrophy varied along the oligotrophic gradient and that nutrient and/or carbon limitation favors BChl-a synthesis.

We investigated the identity of the limiting nutrient of the pelagic microbial food web in the Mediterranean Sea using nutrient manipulated microcosms during summer 2008. Experiments were carried out with surface waters at the center of anticyclonic eddies in the Western Basin, the Ionian Basin, and the Levantine Basin. In situ, the ratio of N to P was always higher in both dissolved and particulate organic fractions compared to the Redfield ratio, suggesting a relative P-starvation. In each experiment, four different treatments in triplicates (addition of ammonium, phosphate, a combination of both, and the unamended control) were employed and chemical and biological parameters monitored throughout a 3–4 day incubation. Temporal changes of turnover time of phosphate and ATP, and alkaline phosphatase activity during the incubation suggested that the phytoplankton and heterotrophic prokaryotes (Hprok) communities were not P-limited at the sites. Furthermore, statistical comparison among treatments at the end of the incubation did not support a hypothesis of P-limitation at the three study sites. In contrast, primary production was consistently limited by N, and Hprok growth was not limited by N nor P in the Western Basin, but N-limited in the Ionian Basin, and N and P co-limited in the Levantine Basin. Our results demonstrated the gap between biogeochemical features (an apparent P-starved status) and biological responses (no apparent P-limitation). We question the general notion that Mediterranean surface waters are limited by P alone during the stratified period.

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

Changes in concentration and composition of Iatroscan-measured dissolved lipids were examined at a daily to month scale, in relation to the hydrological and biological context at a central site of the Ligurian sea, NW Mediterranean during the PECHE-DYNAPROC 2 experiment (14 September to 17 October 2004). Lipid concentrations (excluding hydrocarbons) (TLd-HC) and TLd-HC to DOC ratios in the 0–1000 m water column, varied from 5.3 to 48.5 μg l−1 and 0.01 to 0.09, respectively. The highest TLd-HC concentration values were found in the 0–50 m surface layer, coinciding with phytoplankton biomass. Significant correlations (pn=87) between glycolipids from chloroplast membranes, namely the monogalactosyldiacylglycerols, a major component of dissolved lipids (25.1±10.8% of TLd-HC, n=166), and various phytoplankton pigments (chlorophyll cs-170, violaxanthin, diadinoxanthin, zeaxanthin, and lutein), suggested that picoeucaryote phytoplankton were a major source of dissolved lipids. Lipid metabolites (free fatty acids, alcohols, diacylglycerols and monoacylglycerols), an other important component of TLd-HC (37.6±11.1%, n=166), showed a greater degree of degradation of lipids in this transitional period than previously observed earlier in the year. Zooplankton wax and steryl ester biomarkers (WSE) and triacylglycerols showed a distinct periodicity in the mesopelagic layer throughout the period investigated. Concentrations of WSE (5.5–13.6 μg/l) increased in the 0–150 m surface layer, mid-way through the cruise (4–6 October), before the winter mixing. WSE were observed later and deeper in the mesopelagic layer (6–11 October), accompanied by rebounds in hydrocarbons (6–8 October) and phospholipid concentrations (12 October) in the 400–1000 m depth layer. Zooplankton migration and/or fecal pellet egestion, followed by DOM release from POM, were likely responsible for the appearance of these lipid signatures in the mesopelagic layer. Because we observed these signatures during low wind period only (

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.