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

The tropical Atlantic has been facing a massive proliferation of Sargassum since 2011, with severe environmental and socioeconomic impacts. The development of large-scale modeling of Sargassum transport and physiology is essential to clarify the link between Sargassum distribution and environmental conditions, and to lay the groundwork for a seasonal forecast at the scale of the tropical Atlantic basin. We developed a modeling framework based on the Nucleus for European Modelling of the Ocean (NEMO) ocean model, which integrates transport by currents and waves, and physiology of Sargassum with varying internal nutrients quota, and considers stranding at the coast. The model is initialized from basin-scale satellite observations, and performance was assessed over the year 2017. Model parameters are calibrated through the analysis of a large ensemble of simulations, and the sensitivity to forcing fields like riverine nutrient inputs, atmospheric deposition, and waves is discussed. Overall, results demonstrate the ability of the model to reproduce and forecast the seasonal cycle and large-scale distribution ofSargassum biomass.

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.

A carbonate chemistry balance module was implemented into a biogeochemical model of the planktonic food web. The model, named Eco3M-CarbOx, includes 22 state variables that are dispatched into 5 compartments: phytoplankton, heterotrophic bacteria, detrital particulate organic matter, labile dissolved organic, and inorganic matter. This model is applied to and evaluated in the Bay of Marseille (BoM, France), which is a coastal zone impacted by the urbanized and industrialized Aix–Marseille Metropolis, and subject to significant increases in anthropogenic emissions of CO2.The model was evaluated over the year 2017, for which in situ data of the carbonate system are available in the study site. The biogeochemical state variables of the model only change with time, to represent the time evolution of a sea surface water cell in response to the implemented realistic forcing conditions. The model correctly simulates the value ranges and seasonal dynamics of most of the variables of the carbonate system except for the total alkalinity. Several numerical experiments were conducted to test the response of carbonate system to (i) a seawater temperature increase, (ii) wind events, (iii) Rhône River plume intrusions, and (iv) different levels of atmospheric CO2 contents. This set of numerical experiments shows that the Eco3M-CarbOx model provides expected responses in the alteration of the marine carbonate balance regarding each of the considered perturbation. When the seawater temperature changes quickly, the behavior of the BoM waters alters within a few days from a source of CO2 to the atmosphere to a sink into the ocean. Moreover, the higher the wind speed is, the higher the air–sea CO2 gas exchange fluxes are. The river intrusions with nitrate supplies lead to a decrease in the pCO2 value, favoring the conditions of a sink for atmosphericCO2 into the BoM. A scenario of high atmospheric concentrations of CO2 also favors the conditions of a sink for atmosphericCO2 into the waters of the BoM. Thus the model results suggest that external forcings have an important impact on the carbonate equilibrium in this coastal area.

The Bay of Marseille (BoM), located in the northwestern Mediterranean Sea, is affected by various hydrodynamic processes (e.g., Rhône River intrusion and upwelling events) that result in a highly complex local carbonate system. In any complex environment, the use of models is advantageous since it allows us to identify the different environmental forcings, thereby facilitating a better understanding. By combining approaches from two biogeochemical ocean models and improving the formulation of total alkalinity, we develop a more realistic representation of the carbonate system variables at high temporal resolution, which enables us to study air–sea CO2 fluxes and seawater pCO2 variations more reliably. We apply this new formulation to two particular scenarios that are typical for the BoM: (i) summer upwelling and (ii) Rhône River intrusion events. In both scenarios, our model was able to correctly reproduce the observed patterns of pCO2 variability. Summer upwelling events are typically associated with a pCO2 decrease that mainly results from decreasing near-surface temperatures. Furthermore, Rhône River intrusion events are typically associated with a pCO2 decrease, although, in this case, the pCO2 decrease results from a decrease in salinity and an overall increase in total alkalinity. While we were able to correctly represent the daily range of air–sea CO2 fluxes, the present configuration of Eco3M_MIX-CarbOx does not allow us to correctly reproduce the annual cycle of air–sea CO2 fluxes observed in the area. This pattern directly impacts our estimates of the overall yearly air–sea CO2 flux as, even if the model clearly identifies the bay as a CO2 sink, its magnitude was underestimated, which may be an indication of the limitations inherent in dimensionless models for representing air–sea CO2 fluxes.

A mechanistic physiological model of the appendicularian Oikopleura dioica has been built to represent its three feeding processes (filtration, ingestion and assimilation). The mathematical formulation of these processes is based on laboratory observations from the literature, and tests different hypotheses. This model accounts for house formation dynamics, the food storage capacity of the house and the gut throughput dynamics. The half-saturation coefficient for ingestion resulting from model simulations is approximately 28 [Formula: see text] and is independent of the weight of the organism. The maximum food intake for ingestion is also a property of the model and depends on the weight of the organism. Both are in accordance with data from the literature. The model also provides a realistic representation of carbon accumulation within the house. The modelled half-saturation coefficient for assimilation is approximately 15 [Formula: see text] and is also independent of the weight of the organism. Modelled gut throughput dynamics are based on faecal pellet formation by gut compaction. Model outputs showed that below a food concentration of 30 [Formula: see text], the faecal pellet weight should represent a lower proportion of the body weight of the organism, meaning that the faecal pellet formation is not driven by gut filling. Simulations using fluctuating environmental food availability show that food depletion is not immediately experienced by the organism but that it occurs after a lag time because of house and gut buffering abilities. This lag time duration lasts at least 30 minutes and can reach more than 2 hours, depending on when the food depletion occurs during the house lifespan.

Many current biogeochemical models rely on an autotrophic versus heterotrophic food web representation. However, in recent years, an increasing number of studies have begun to challenge this approach. Several authors have highlighted the importance of protists capable of combining photoautotrophic and heterotrophic nutrition in a single cell. These mixotrophic protists are known to play an important role in the carbon cycle. Here, we present a new biogeochemical model that represents the food web using variable stoichiometry. It contains the classic compartments such as zooplankton, phytoplankton, and heterotrophic bacteria and a newly added compartment to represent two types of mixotrophic protists: non-constitutive mixotrophs (NCMs) and constitutive mixotrophs (CMs). We demonstrate that the model correctly reproduces the characteristics of NCMs and CMs and proceed to study the impact of light and nutrient limitation on planktonic ecosystem structure in a highly dynamic Mediterranean coastal area, namely the Bay of Marseille (BoM, France), paying special attention to the dynamics of mixotrophic protists in these limiting conditions. In addition, we investigate the carbon, nitrogen, and phosphorus fluxes associated with mixotrophic protists and showed the following: (i) the portion of the ecosystem in terms of the percentage of carbon biomass occupied by NCMs decreases when resources (nutrient and prey concentrations) decrease, although their mixotrophy allows them to maintain a carbon biomass almost as significant as the copepod one (129.8 and 148.7 mmolCm-3, respectively), as photosynthesis increases as a food source, and (ii) the portion of the ecosystem in terms of the percentage of carbon biomass occupied by CM increases when nutrient concentrations decrease due to their capability to ingest prey to supplement their N and P needs. In addition to providing new insights regarding the conditions that lead to the emergence of mixotrophs in the BoM, this work provides a new tool to perform long-term studies and predictions of mixotroph dynamics in coastal environments under different environmental forcings.

Since 2004, the Service facility SNAPO-CO2 (Service National d’Analyse des Paramètres Océaniques du CO2) housed by the LOCEAN laboratory (Paris, France) has been in charge for the analysis of Total Alkalinity (AT) and Total dissolved inorganic carbon (CT) of seawater samples on a series of cruises or ships of opportunity conducted in different regions in the frame of French projects. More than 44000 observations are synthetized in this work. Sampling was performed either from CTD-Rosette casts (Niskin bottles) or collected from the ship’s seawater supply (intake at about 5m depth). After completion of each cruise, discrete samples were returned back at LOCEAN laboratory and stored in a dark room at 4 °C before analysis generally within 2-3 months after sampling (sometimes within a week). AT and CT were analyzed simultaneously by potentiometric titration using a closed cell (Edmond, 1970). Certified Reference Materials (CRMs) provided by Pr. A. Dickson (Scripps Institution of Oceanography, San Diego, USA) were used to calibrate the measurements. The same instrumentation was used for underway measurements during OISO cruises (https://doi.org/10.18142/228) and OISO AT-CT data for 1998-2018 in the South Indian Ocean added in this synthesis. The synthesis is organized in two files (one for Global ocean and the Coastal Zones, one for the Mediterranean Sea) with the same format: Cruise name, Ship name, day, month, year, hour, minute, second, latitude, longitude, depth, AT (µmol/kg), Flag-AT, CT (µmol/kg), Flag-CT, Temperature (°C), Flag-Temp, Salinity (PSU), Flag-Salinity, nsample/cruise, nsample on file, sampling method.

Evidence of phosphorus limitation of algal C- and N-uptake in a NW Mediterranean coastal area (Gulf of Lions) was obtained from a field survey of inorganic and organic N, P and C and from bioassays carried out during the late winter-early spring 1998. Dissolved inorganic nitrogen (ΣDIN = NO₃ + NO₂ + NH₄) and phosphorus (DIP) distributions showed a clear DIP depletion in the inorganic fractions available for primary production. While below the 150 m depth, the mean ΣDIN to DIP ratio was close to the typical Mediterranean ratio of 22, while values found in the upper layer (0 to 150 m) were about 3 times higher (68.4:1 on average). In this upper layer, N:P (19.9:1) and C:P (159.7:1) ratios in the particulate organic matter were higher than the Redfield ratio and also indicated P depletion in this fraction. In the dissolved organic pool, P depletion was higher than in the particulate organic pool, since the mean C:N:P ratios were 1674:75:1 in the photic layer. Dissolved organic forms of C and N represented the bulk (ca 94 and 86%, respectively) of the total organic matter, while ca 31% of the organic P was in particulate fraction. The apparent imbalance between N and P in the inorganic fraction was partly attributed to an imbalance in the corresponding nutrient utilization by the phytoplanktonic community, and partly due to the influence of the Rhone River. Additions of small amounts of DIP to surface samples led (1) to a decrease in C-uptake (≅30%) during the first 24 h incubation, (2) to a rapid increase in chlorophyll biomass and (3) to stimulate nitrate uptake (≅60%), suggesting DIP limitation of new production and of algal biomass during the spring 1998 in the Gulf of Lions.

Nitrogen regeneration fluxes of ammonium (NH4+) and nitrate (NO3–) as well as losses of dissolved organic nitrogen (DON) by phytoplankton were investigated over a 2 mo period (spring 1997) in a NW coastal Mediterranean area (Gulf of Lions) using 15N-tracer techniques. Profiles of dissolved inorganic nitrogen (DIN) concentrations were almost uniform with values of 600, 150 and 35 nM for NO3–, NO2– and NH4+, respectively, except at the end of the study period when the upper layer became nitrogen-depleted (<50 nM down to 40 m). Chlorophyll (chl) distributions showed a surface maximum (to 0.85 mg m–3) and a deep maximum (to 1.25 mg m–3) at 40 m. Plankton DIN utilization (net uptake) was most of the time highest at the surface, with rates reaching 62 and 40 nM d–1 for NH4+ and NO3–, respectively. However, a deepening (to 60 m) of maximum NO3– uptake rates with a corresponding deepening of the nitracline sometimes occurred during the experiment. Therefore, f-ratio profiles depicted maximum surface values (∼0.40) at the beginning of the experiment and a deep maximum at the end. NH4+ regeneration rates were 1 order of magnitude higher (up to 220 nM d–1) than nitrification and DIN loss (as DON) rates, and could largely sustain more than 100% of the plankton NH4+ demand. Underestimation of NH4+ uptake rates due to 15N isotope dilution had only a small effect on the f-ratio calculation (overestimation <5%). Nitrification occurred from the surface (10 to 20 nM d–1) down to the base of the euphotic layer (30 nM d–1), and corresponded to 90% and >>100% of the plankton NO3– demand at the surface and in the nitracline, respectively. Consequently, a great part of NO3– uptake did not correspond to new production and should be considered as regenerated production, particularly in the NO3– depleted surface layer. Profiles of DIN loss (as DON) well paralleled those of DIN net uptake with values highest at surface reaching 35 and 14 nM d–1 for NH4+ and NO3–, respectively. DIN loss rates represented on average ∼23% of gross DIN uptake (gross DIN uptake = DIN losses + DIN net uptake) whatever the substrate was, indicating that (1) DIN loss (as DON) did not depend on the nitrogen source, and (2) DIN uptake was mostly due to phytoplankton and not to bacterioplankton, although the study area tended to be globally nitrogen-depleted and based on regeneration. Failure to account for DIN losses had no significant effect on the computation of f-ratios.