Luca Polimene

Substantial energy input to the mesopelagic ecosystem from the seasonal mixed-layer pump

The “mesopelagic” is the region of the ocean between about 100 and 1000 m that harbours one of the largest ecosystems and fish stocks on the planet1,2. This vastly unexplored ecosystem is believed to be mostly sustained by chemical energy, in the form of fast-sinking particulate organic carbon, supplied by the biological carbon pump3. Yet, this supply appears insufficient to match mesopelagic metabolic demands4–6. The mixed-layer pump is a physically-driven biogeochemical process7–11 that could further contribute to meet these energetic requirements. However, little is known about the magnitude and spatial distribution of this process at the global scale. Here we show that the mixed-layer pump supplies an important seasonal flux of organic carbon to the mesopelagic. By combining mixed-layer depths from Argo floats with satellite retrievals of particulate organic carbon, we estimate that this pump exports a global flux of about 0.3 Pg C yr−1 (range 0.1 – 0.5 Pg C yr−1). In high-latitude regions where mixed-layers are deep, this flux is on average 23%, but can be greater than 100% of the carbon supplied by fast sinking particles. Our results imply that a relatively large flux of organic carbon is missing from current energy budgets of the mesopelagic.

A numerical simulation study of dissolved organic carbon accumulation in the northern Adriatic Sea

The leading author of this paper was supported by a Ph.D. fellowship given to the Environmental Science graduate program of the University of Bologna at Ravenna and by the VECTOR project funded by the Italian Ministry of Research and University. N. Pinardi and M. Zavatarelli were partially supported by the MFSTEP project (EU contract EVK3-CT-2002-00075) and the ADRICOSM Project (funded by the Italian Ministry of Environment and Territory, Division of Environmental Research and Development). Icarus Allen and Marcello Vichi acknowledge the support by the EUR-OCEANS network of excellence (contract 511106).

Modelling the Stoichiometric Regulation of C-Rich Toxins in Marine Dinoflagellates

Toxin production in marine microalgae was previously shown to be tightly coupled with cellular stoichiometry. The highest values of cellular toxin are in fact mainly associated with a high carbon to nutrient cellular ratio. In particular, the cellular accumulation of C-rich toxins (i.e., with C:N > 6.6) can be stimulated by both N and P deficiency. Dinoflagellates are the main producers of C-rich toxins and may represent a serious threat for human health and the marine ecosystem. As such, the development of a numerical model able to predict how toxin production is stimulated by nutrient supply/deficiency is of primary utility for both scientific and management purposes. In this work we have developed a mechanistic model describing the stoichiometric regulation of C-rich toxins in marine dinoflagellates. To this purpose, a new formulation describing toxin production and fate was embedded in the European Regional Seas Ecosystem Model (ERSEM), here simplified to describe a monospecific batch culture. Toxin production was assumed to be composed by two distinct additive terms; the first is a constant fraction of algal production and is assumed to take place at any physiological conditions. The second term is assumed to be dependent on algal biomass and to be stimulated by internal nutrient deficiency. By using these assumptions, the model reproduced the concentrations and temporal evolution of toxins observed in cultures of Ostreopsis cf. ovata, a benthic/epiphytic dinoflagellate producing C-rich toxins named ovatoxins. The analysis of simulations and their comparison with experimental data provided a conceptual model linking toxin production and nutritional status in this species. The model was also qualitatively validated by using independent literature data, and the results indicate that our formulation can be also used to simulate toxin dynamics in other dinoflagellates. Our model represents an important step towards the simulation and prediction of marine algal toxicity.

Impacts of light shading and nutrient enrichment geo-engineering approaches on the productivity of a stratified, oligotrophic ocean ecosystem

Geo-engineering proposals to mitigate global warming have focused either on methods of carbon dioxide removal, particularly nutrient fertilization of plant growth, or on cooling the Earths surface by reducing incoming solar radiation (shading). Marine phytoplankton contribute half the Earths biological carbon fixation and carbon export in the ocean is modulated by the actions of microbes and grazing communities in recycling nutrients. Both nutrients and light are essential for photosynthesis, so understanding the relative influence of both these geo-engineering approaches on ocean ecosystem production and processes is critical to the evaluation of their effectiveness. In this paper, we investigate the relationship between light and nutrient availability on productivity in a stratified, oligotrophic subtropical ocean ecosystem using a one-dimensional water column model coupled to a multi-plankton ecosystem model, with the goal of elucidating potential impacts of these geo-engineering approaches on ecosystem production. We find that solar shading approaches can redistribute productivity in the water column but do not change total production. Macronutrient enrichment is able to enhance the export of carbon, although heterotrophic recycling reduces the efficiency of carbon export substantially over time. Our results highlight the requirement for a fuller consideration of marine ecosystem interactions and feedbacks, beyond simply the stimulation of surface blooms, in the evaluation of putative geo-engineering approaches.

Biological or microbial carbon pump? The role of phytoplankton stoichiometry in ocean carbon sequestration

Once fixed by photosynthesis carbon becomes part of the marine food web. The fate of this carbon has two possible outcomes: it may be respired and released back to the ocean and potentially to the atmosphere as CO2 or retained in the ocean interior and/or marine sediments for extended time scales. The most important biologically mediated processes responsible for long term carbon storage in the ocean are the biological carbon pump (BCP) and the microbial carbon pump (MCP). While acting simultaneously in the ocean, the balance between these two mechanisms is thought to vary depending on the trophic state of the environment. Using previously published formulations, we propose a modelling framework to simulate variability in the MCP: BCP ratio as a function of external nutrients. Our results suggest that the role of the MCP might become more significant under future climate change conditions where increased stratification enhances the oligotrophic nature of the surface ocean. Based on these model results, we propose a conceptual framework in which the internal stoichiometry of phytoplankton, modulating both grazing pressure and DOM production (via phytoplankton exudation), plays a crucial role in regulating the MCP: BCP ratio.

EFFECTS OF ULTRAVIOLET-A RADIATION AND NUTRIENT AVAILABILITY ON THE CELLULAR COMPOSITION OF PHOTOPROTECTIVE COMPOUNDS IN GLENODINIUM FOLIACEUM (DINOPHYCEAE)(1).

The photoprotective response in the dinoflagellate Glenodinium foliaceum F. Stein exposed to ultraviolet‐A (UVA) radiation (320–400 nm; 1.7 W · m2) and the effect of nitrate and phosphate availability on that response have been studied. Parameters measured over a 14 d growth period in control (PAR) and experimental (PAR + UVA) cultures included cellular mycosporine‐like amino acids (MAAs), chls, carotenoids, and culture growth rates. Although there were no significant effects of UVA on growth rate, there was significant induction of MAA compounds (28 ± 2 pg · cell−1) and a reduction in chl a (9.6 ± 0.1 pg · cell−1) and fucoxanthin (4.4 ± 0.1 pg · cell−1) compared to the control cultures (3 ± 1 pg · cell−1, 13.3 ± 3.2 pg · cell−1, and 7.4 ± 0.3 pg · cell−1, respectively). In a second investigation, MAA concentrations in UVA‐exposed cultures were lower when nitrate was limited (P < 0.05) but were higher when phosphate was limiting. Nitrate limitation led to significant decreases (P < 0.05) in cellular concentration of chls (chl c1, chl c2, and chl a), but other pigments were not affected. Phosphate availability had no effect on final pigment concentrations. Results suggest that nutrient availability significantly affects cellular accumulation of photoprotective compounds in G. foliaceum exposed to UVA.

Assimilation of Ocean‐Color Plankton Functional Types to Improve Marine Ecosystem Simulations

We assimilated plankton functional types (PFTs) derived from ocean colour into a marine ecosystem model, to improve the simulation of biogeochemical indicators and emerging properties in a shelf sea. Error-characterized chlorophyll concentrations of four PFTs (diatoms, dinoflagellates, nanoplankton and picoplankton), as well as total chlorophyll for comparison, were assimilated into a physical-biogeochemical model of the North East Atlantic, applying a localized Ensemble Kalman filter. The reanalysis simulations spanned the years 1998 to 2003. The skill of the reference and reanalysis simulations in estimating ocean colour and in situ biogeochemical data were compared by using robust statistics. The reanalysis outperformed both the reference and the assimilation of total chlorophyll in estimating the ocean-colour PFTs (except nanoplankton), as well as the not-assimilated total chlorophyll, leading the model to simulate better the plankton community structure. Crucially, the reanalysis improved the estimates of not-assimilated in situ data of PFTs, as well as of phosphate and pCO2, impacting the simulation of the air-sea carbon flux. However, the reanalysis increased further the model overestimation of nitrate, in spite of increases in plankton nitrate uptake. The method proposed here is easily adaptable for use with other ecosystem models that simulate PFTs, for, e.g., reanalysis of carbon fluxes in the global ocean and for operational forecasts of biogeochemical indicators in shelf-sea ecosystems.

Contribution of structural recalcitrance to the formation of the deep oceanic dissolved organic carbon reservoir: Structural recalcitrance dominates DOM persistence

The origin of the recalcitrant dissolved organic carbon (RDOC) reservoir in the deep ocean remains enigmatic. The structural recalcitrance hypothesis suggests that RDOC is formed by molecules that are chemically resistant to bacterial degradation. The dilution hypothesis claims that RDOC is formed from a large diversity of labile molecules that escape bacterial utilization due to their low concentrations, termed as RDOCc . To evaluate the relative contributions of these two mechanisms in determining the long-term persistence of RDOC, we model the dynamics of both structurally recalcitrant DOC and RDOCc based on previously published data that describes deep oceanic DOC degradation experiments. Our results demonstrate that the majority of DOC (84.5 ± 2.2%) in the deep ocean is structurally recalcitrant. The intrinsically labile DOC (i.e., labile DOC that rapidly consumed and RDOCc ) accounts for a relatively small proportion and is consumed rapidly in the incubation experiments, in which 47.8 ± 3.2% of labile DOC and 21.9 ± 4.6% of RDOCc are consumed in 40 days. Our results suggest that the recalcitrance of RDOC is largely related to its chemical properties, whereas dilution plays a minor role in determining the persistence of deep-ocean DOC.

A substantial fraction of phytoplankton-derived DON is resistant to degradation by a metabolically versatile, widely distributed marine bacterium

The capacity of bacteria for degrading dissolved organic nitrogen (DON) and remineralising ammonium is of importance for marine ecosystems, as nitrogen availability frequently limits productivity. Here, we assess the capacity of a widely distributed and metabolically versatile marine bacterium to degrade phytoplankton-derived dissolved organic carbon (DOC) and nitrogen. To achieve this, we lysed exponentially growing diatoms and used the derived dissolved organic matter (DOM) to support an axenic culture of Alteromonas sp.. Bacterial biomass (as particulate carbon and nitrogen) was monitored for 70 days while growth dynamics (cell count), DOM (DOC, DON) and dissolved nutrient concentrations were monitored for up to 208 days. Bacterial biomass increased rapidly within the first 7 days prior to a period of growth/death cycles potentially linked to rapid nutrient recycling. We found that ≈75% of the initial DOC and ≈35% of the initial DON were consumed by bacteria within 40 and 4 days respectively, leaving a significant fraction of DOM resilient to degradation by this bacterial species. The different rates and extents to which DOC and DON were accessed resulted in changes in DOM stoichiometry and the iterative relationship between DOM quality and bacterial growth over time influenced bacterial cell C:N molar ratio. C:N values increased to 10 during the growth phase before decreasing to values of ≈5, indicating a change from relative N-limitation/C-sufficiency to relative C-limitation/N-sufficiency. Consequently, despite its reported metabolic versatility, we demonstrate that Alteromonas sp. was unable to access all phytoplankton derived DOM and that a bacterial community is likely to be required. By making the relatively simple assumption that an experimentally derived fraction of DOM remains resilient to bacterial degradation, these experimental results were corroborated by numerical simulations using a previously published model describing the interaction between DOM and bacteria in marine systems, thus supporting our hypothesis.