Philippe Souchu

Modelling nitrogen, primary production and oxygen in a Mediterranean lagoon. Impact of oysters farming and inputs from the watershed

An ecosystem model based on nitrogen cycling and oxygen has been developed for the Thau lagoon. It takes into account the specific features of this Mediterranean lagoon, a semi-confined system with watershed inputs and oyster farming. The ecosystem model uses currents calculated by a two-dimensional hydrodynamic model and integrated into a box model. This model is compared with a year survey data and used to estimate nitrogen and oxygen fluxes between the different ecosystem compartments. The yearly simulation shows that the ecosystem behavior is driven by meteorological forcing, especially rain which causes watershed inputs. These inputs trigger microphytoplankton growth, which is responsible for new primary production. During dry periods, nitrogen is recycled into the lagoon thanks to oysters excretion, sediment release, microzooplankton excretion and mineralization. Ammonium produced in this way is consumed by a population of pico- and nanophytoplankton causing regenerated primary production. Consequently, the ecosystem remains highly productive in summer even without external inputs. Shellfish farming also plays an important role in the whole lagoon through biodeposition. Driven by biodeposition, sediment release is the major source of nitrogen in the water column and causes oxygen reduction. The oysters contribute to the recycling activity by excretion, which supports the regenerated primary production. They are also involved in oxygen consumption by respiration which can cause local hypoxia. Further improvements are proposed before this model may become a functional environmental model for a lagoon ecosystem.

NITROGENOUS NUTRITION OF ALEXANDRIUM CATENELLA (DINOPHYCEAE) IN CULTURES AND IN THAU LAGOON, SOUTHERN FRANCE1

Alexandrium catenella (Whedon et Kofoid) Balech was isolated from Thau lagoon (northern Mediterranean) and its growth and uptake characteristics measured for nitrate, ammonium, and urea. Although affinity constants did not indicate a preference for ammonium over nitrate, there was a strong inhibition of nitrate uptake by ammonium when both nitrogen (N) sources were present. Nitrogen budgets during growth in cultures revealed major imbalances between decreases in dissolved N and increases in particulate N, indicating excretion of dissolved organic N during the early part of the growth phase and uptake during the later part. A quasi‐unialgal bloom in November 2001 (4×106 cells·L−1) allowed measurements of uptake of nitrate, nitrite, ammonium, and urea; net and gross growth rate of A. catenella; and grazing rates on this organism. The affinity constants indicate that it is not a strong competitor for the N nutrients tested when these are in low concentrations (<10 μgat N·L−1), compared with other members of the phytoplankton community. Indirect evidence from cultures indicate that dissolved organic N compounds could be important in triggering those blooms. Finally, the strongly unbalanced growth observed in the field indicates that A. catenella exhibits a storage rather than a growth response to a nutrient pulse and is adapted to low frequency events such as the passage of frontal disturbances. The disappearance of A. catenella was due to grazing that balanced growth at the peak of the bloom.

ACCLIMATION OF NITRATE UPTAKE BY PHYTOPLANKTON TO HIGH SUBSTRATE LEVELS

A literature review of data on nitrate uptake by phytoplankton suggests that nitrate levels above 20 μmol N·L−1 generally stimulated uptake rates in cultured unicellular algae and natural phytoplankton communities. This phenomenon indicates that phytoplankton cells acclimate to elevated nitrate levels by increasing their uptake capacity in a range of concentrations previously considered to be saturating. Cyanobacteria and flagellates were found to present a considerable capacity for acclimation, with low (0.1–2 μmol N·L−1) half‐saturation values (Ks) at low (5–20 μmol N·L−1) substrate levels and high (1–80 μmol N·L−1) Ks values at high (30–100 μmol N·L−1) substrate levels. However, some diatom genera (Rhizosolenia, Skeletonema, Thalassiosira) also appeared to possess a low affinity nitrate uptake system (Ks between 18 and 120 μmol N·L−1), which can help resolve the paradox of their presence in enriched seas. It follows that present models of nitrate uptake can severely underestimate the effects of high nitrate concentrations on phytoplankton dynamics and development. A more adequate approach would be to consider the possibility of multiphasic uptake involving several phase transitions as nitrate concentrations increased. Because it is a nonlinear phenomenon featuring strong thresholds, this effect appears to override that of other variables, such as irradiance, temperature, and cell size. Within the present context of eutrophication and for a range of concentrations that is becoming more and more ecologically relevant, equations are tentatively presented as a first approach to estimate Ks from ambient nitrate concentrations.

Relations milieu-ressources · impact de la conchyliculture sur un environnement lagunaire méditerranéen (Thau)

Abstract Shellfish farming leaves its mark on the environment in which it has developed, and the men who depend upon it. These changes have altogether balanced the lagoon cycle and have caused disastrous episodic events. Increased water clarity caused by the uptake of particulate material by shellfish farming allows seagrass to grow in deeper areas of the lagoon (down to five metres). Shellfish farming nutrient transformations increase ecosystem productivity, even if the filtration pressure keeps phytoplankton biomass at a low level. Storage of phosphorus and nitrogen in animal tissue limits eutrophication in this ecosystem. Transfer of oysters from growout facilities increases animal and vegetal specific diversity. The presence of large amounts of shellfish allows for the development of a masive benthos, while organic enrichment from biodeposition changes the specific composition of soft-bottom benthos. In the deeper areas, (less than six metres), where summer thermoclines limit oxygen transfer from surface water, the organically enriched substrate induces oxygen depletion and ammonium and nitrogen sulfide accumulation in the water column. This ecosystem dysfunction kills benthic populations, and sometimes reaches pelagic populations and affects the shellfish farming economy.

Patterns in nutrient limitation and chlorophyll a along an anthropogenic eutrophication gradient in French Mediterranean coastal lagoons

A cross-ecosystem comparison of data obtained from 20 French Mediterranean lagoons with contrasting eutrophication status provided the basis for investigating the variables that best predict chlorophyll a (Chl a) concentrations and nutrient limitation of phytoplankton biomass along a strong nutrient enrichment gradient. Summer concentrations of dissolved inorganic nitrogen (DIN) and phosphorus (DIP) comprised only a small fraction of total nitrogen (TN) and total phosphorus (TP). On the basis of inorganic nutrient concentrations, the most oligotrophic lagoons appeared to be phosphorus-limited, with a tendency towards the development of nitrogen limitation as eutrophication increased, as evidenced by decreasing DIN:DIP ratios. A weak but significantly positive relationship was found between dissolved silicate (DSi) and Chl a, reflecting DSi accumulation in the water column along the trophic state gradient and implying a progressive shift away from potential Si limitation of phytoplankton growth. Observed concentrations of Chl a were far better explained by TN and TP than by DIN and DIP concentrations, suggesting that a total nutrient based approach is likely to be the most appropriate for managing eutrophication in Mediterranean lagoons and other coastal waters. These results give credence to the idea that marine and freshwater environments respond in a similar fashion to nutrient enrichment.

Modélisation numérique des crises anoxiques (malaïgues) dans la lagune de Thau (France)

Abstract In summer, the Thau lagoon is often affected by anoxic outbreaks, which induce shellfish mortality. A 3D hydrodynamic model has been set up to simulate currents driven by the wind and the sea temperature. It has been coupled to a biochemical model including oxygen, organic matter and sulphides to reproduce the ‘malaigue’ event. Firstly the model has been applied to the 1997 anoxic crisis that broke out in the Meze area. It raises the importance of wind data and the impact of shellfish farming structures that lower velocity currents. The model has also been applied to the 1982 and 1987 anoxic events, using historical data. The results point out that the ‘malaigue’ depends on the localization of the original flare up spots and the trophic status of the lagoon. Secondly, the model has been used to assess different modifications on locations of shellfish farming areas in order to find the best way of reducing the anoxic crisis. The oxygen level could be improved by extending the distance between the shellfish areas and the coast or by reducing shellfish density.

Response of coastal phytoplankton to ammonium and nitrate pulses: seasonal variations of nitrogen uptake and regeneration

Seasonal variation in uptake and regeneration of ammonium and nitrate in a coastal lagoon was studied using 15N incorporation in particulate matter and by measuring changes in particulate nitrogen. Uptake and regeneration rates were two orders of magnitude lower in winter than in summer. Summer uptake values were 2.8 and 2.2 μmol N.l−1.d−1 for ammonium and nitrate, respectively. Regeneration rates were 2.9 and 2.1 μmol N.l−1.d−1 for ammonium and nitrate respectively. Regeneration/uptake ratios were often below one, indicating that water column processes were not sufficient to satisfy the phytoplankton nitrogen demand. This implies a role of other sources of nitrogen, such as macrofauna (oysters and epibionts) and sediment. Phytoplankton was well adapted to the seasonal variations in resources, with mixotrophic dinoflagellates dominant in winter, and fast growing diatoms in summer. In winter and spring, ammonium was clearly preferred to nitrate as a nitrogen source, but nitrate was an important nitrogen source in summer because of high nitrification rates. Despite low nutrient levels, the high rates of nitrogen regeneration in summer as well as the simultaneous uptake of nitrate and ammonium allow high phytoplankton growth rates which in turn enable high oyster production.

Oxygen fluxes involving the benthic micro- and macrophytic components in the Thau Lagoon under pre-anoxic conditions

Measurements of oxygen fluxes at the water-sediment interface (transparent and opaque enclosures) were made on bare sediments inhabited by microphytes on the one hand, and on sediments diversely colonized by macrophytes (macroalgae and seagrasses) on the other hand. Five stations, typical of different biotopes of a Mediterranean shallow lagoon were studied from May to July 1993 in order to observe an anoxic event which usually occurs at that time of the year. Average diurnal respiration of benthic communities ranged from 40 mg (or 1.25 mmol) O2 m2 h−1 in bare sediments (31 % of the lagoon area) to 76 mg (or 2.37 mmol) O2 m−2 h−1 in sediments with a medium coverage of macroalgae (37 % of the total area) and, finally, to 100 mg (or 3.12 mmol) O2 m−2h−1 in the denser macrophytic area (32 % of the total area). The highest diurnal gross production was observed in the zone colonized by macroalgae and seagrasses, and especially in corridors between shellfish-cultivation tables (300 mg O2 m−2 h−1, or 9.37 mmol O2m−2 h−1, equivalent to 113 mg C m−2 h−1). Overall, during this period, net bottom oxygen production was close to nil in ca. 30 %, and positive in 70 % of the lagoon area. The average net oxygen production for the whole lagoon in summer is thus in the order of 100 mg O2 m−2h−1. In 1993, at the end of July, an anoxic event was avoided due to a period of strong wind.

Oyster Farming Impact on the Environment of a Mediterranean Lagoon (THAU)(Preliminary results of the OXYTHAU programme)

The lagoon of Thau is a shallow microtidal system exposed to terrestrial and marine influences. The residence time and the vertical mixing of water-mass are controlled by winds. Periodical absence of wind in Summer can lead to strong anoxia in sediments and bottom waters. Moreover, low inorganic nitrogen and chlorophyll-a levels confer seasonal oligotrophic features to the ecosystem. Nevertheless, a large productive stock of oysters (Crassostrea gigas) cultivated on suspended lines and colonized by epibiota (invertebrates, algae, etc...) covers 20 % of the total area. The aim of the present programme (OXYTHAU 1991–1995) is to study effects of oyster farming on the environment of the lagoon of Thau. Preliminary results obtained are presented here.

Influence of nitrogen enrichment on size-fractionated in vitro carboxylase activities of phytoplankton from Thau Lagoon (Coastal Mediterranean Lagoon, France)

The influence of dissolved inorganic and organic nitrogen (DIN and DON) enrichments on pools of enzymes responsible for CO2 fixation by the Calvin-Benson (Rubisco) and beta-carboxylation pathways (beta-carboxylases) were studied in a natural plankton assemblage. The plankton community from a coastal Mediterranean lagoon were incubated in situ for 24 h with initially ammonium, nitrate and DON (taurine) enrichments and compared to a control without any enrichment. An increase of small picophytoplankton and diatom biomass was observed in the enriched inorganic nitrogen treatments 7 - 10 h after the initial enrichment. Phytoplankton biomass decreased in the control and under taurine enrichment suggesting an inorganic nitrogen limitation of phytoplankton growth.