Modelling the combined impacts of climate change and direct anthropogenic drivers on the ecosystem of the northwest European continental shelf

The potential response of the marine ecosystem of the northwest European continental shelf to climate change under a medium emissions scenario (SRES A1B) is investigated using the coupled hydrodynamics-ecosystem model POLCOMS-ERSEM. Changes in the near future (2030–2040) and the far future (2082–2099) are compared to the recent past (1983–2000). The sensitivity of the ecosystem to potential changes in multiple anthropogenic drivers (river nutrient loads and benthic trawling) in the near future is compared to the impact of changes in climate. With the exception of the biomass of benthic organisms, the influence of the anthropogenic drivers only exceeds the impact of climate change in coastal regions. Increasing river nitrogen loads has a limited impact on the ecosystem whilst reducing river nitrogen and phosphate concentrations affects net primary production(netPP) and phytoplankton and zooplankton biomass. Direct anthropogenic forcing is seen to mitigate/amplify the effects of climate change. Increasing river nitrogen has the potential to amplify the effects of climate change at the coast by increasing netPP. Reducing river nitrogen and phosphate mitigates the effects of climate change for netPP and the biomass of small phytoplankton and large zooplankton species but amplifies changes in the biomass of large phytoplankton and small zooplankton.

Environmental selection and resource allocation determine spatial patterns in picophytoplankton cell size

Here we describe a new trait-based model for cellular resource allocation that we use to investigate the relative importance of different drivers for small cell size in phytoplankton. Using the model, we show that increased investment in nonscalable structural components with decreasing cell size leads to a trade-off between cell size, nutrient and light affinity, and growth rate. Within the most extreme nutrient-limited, stratified environments, resource competition theory then predicts a trend toward larger minimum cell size with increasing depth. We demonstrate that this explains observed trends using a marine ecosystem model that represents selection and adaptation of a diverse community defined by traits for cell size and subcellular resource allocation. This framework for linking cellular physiology to environmental selection can be used to investigate the adaptive response of the marine microbial community to environmental conditions and the adaptive value of variations in cellular physiology.

Modelling a light-driven phytoplankton succession

We used a numerical model to investigate if and to what extent cellular photoprotective capacity accounts for succession and vertical distribution of marine phytoplankton species/groups. A model describing xanthophyll photoprotective activity in phytoplankton has been implemented in the European Regional Sea Ecosystem Model and applied at the station L4 in the Western English Channel. Primary producers were subdivided into three phytoplankton functional types defined in terms of their capacity to acclimate to different light-specific environments: low light (LL-type), high light (HL-type) and variable light (VL-type) adapted species. The LL-type is assumed to have low cellular level of xanthophyll-cycling pigments (PX) relative to the modelled photosynthetically active pigments (chlorophyll and fucoxanthin (FUCO) = PSP). The HL-type has high PX content relative to PSP while VL-type presents an intermediate PX to PSP ratio. Furthermore, the VL-type is capable of reversibly converting FUCO to PX and synthesizing new PX under high-light stress. In order to reproduce phytoplankton community succession with each of the three groups being dominant in different periods of the year, we had also to assume reduced grazing pressure on HL-adapted species. Model simulations realistically reproduce the observed seasonal patterns of pigments and nutrients highlighting the reasonability of the underpinning assumptions. Our model suggests that pigment-mediated photophysiology plays a primary role in determining the evolution of marine phytoplankton communities in the winter-spring period corresponding to the shoaling of the mixed layer and the increase of light intensity. Grazing selectivity however contributes to the phytoplankton community composition in summer.

Multiple environmental controls on phytoplankton growth strategies determine adaptive responses of the N : P ratio

The controls on the 'Redfield' N:P stoichiometry of marine phytoplankton and hence the N:P ratio of the deep ocean remain incompletely understood. Here, we use a model for phytoplankton ecophysiology and growth, based on functional traits and resource-allocation trade-offs, to show how environmental filtering, biotic interactions, and element cycling in a global ecosystem model determine phytoplankton biogeography, growth strategies and macromolecular composition. Emergent growth strategies capture major observed patterns in marine biomes. Using a new synthesis of experimental RNA and protein measurements to constrain per-ribosome translation rates, we determine a spatially variable lower limit on adaptive rRNA:protein allocation and hence on the relationship between the largest cellular P and N pools. Comparison with the lowest observed phytoplankton N:P ratios and N:P export fluxes in the Southern Ocean suggests that additional contributions from phospholipid and phosphorus storage compounds play a fundamental role in determining the marine biogeochemical cycling of these elements.

Assimilation of remotely-sensed optical properties to improve marine biogeochemistry modelling

In this paper we evaluate whether the assimilation of remotely-sensed optical data into a marine ecosystem model improves the simulation of biogeochemistry in a shelf sea. A localized Ensemble Kalman filter was used to assimilate weekly diffuse light attenuation coefficient data, Kd(443) from SeaWiFs, into an ecosystem model of the western English Channel. The spatial distributions of (unassimilated) surface chlorophyll from satellite, and a multivariate time series of eighteen biogeochemical and optical variables measured in situ at one long-term monitoring site were used to evaluate the system performance for the year 2006. Assimilation reduced the root mean square error and improved the correlation with the assimilated Kd(443) observations, for both the analysis and, to a lesser extent, the forecast estimates, when compared to the reference model simulation. Improvements in the simulation of (unassimilated) ocean colour chlorophyll were less evident, and in some parts of the Channel the simulation of this data deteriorated. The estimation errors for the (unassimilated) in situ data were reduced for most variables with some exceptions, e.g. dissolved nitrogen. Importantly, the assimilation adjusted the balance of ecosystem processes by shifting the simulated food web towards the microbial loop, thus improving the estimation of some properties, e.g. total particulate carbon. Assimilation of Kd(443) outperformed a comparative chlorophyll assimilation experiment, in both the estimation of ocean colour data and in the simulation of independent in situ data. These results are related to relatively low error in Kd(443) data, and because it is a bulk optical property of marine ecosystems. Assimilation of remotely-sensed optical properties is a promising approach to improve the simulation of biogeochemical and optical variables that are relevant for ecosystem functioning and climate change studies.

Challenges in integrative approaches to modelling the marine ecosystems of the North Atlantic: Physics to fish and coasts to ocean

It has long been recognised that there are strong interactions and feedbacks between climate, upper ocean biogeochemistry and marine food webs, and also that food web structure and phytoplankton community distribution are important determinants of variability in carbon production and export from the euphotic zone. Numerical models provide a vital tool to explore these interactions, given their capability to investigate multiple connected components of the system and the sensitivity to multiple drivers, including potential future conditions. A major driver for ecosystem model development is the demand for quantitative tools to support ecosystem-based management initiatives. The purpose of this paper is to review approaches to the modelling of marine ecosystems with a focus on the North Atlantic Ocean and its adjacent shelf seas, and to highlight the challenges they face and suggest ways forward. We consider the state of the art in simulating oceans and shelf sea physics, planktonic and higher trophic level ecosystems, and look towards building an integrative approach with these existing tools. We note how the different approaches have evolved historically and that many of the previous obstacles to harmonisation may no longer be present. We illustrate this with examples from the on-going and planned modelling effort in the Integrative Modelling Work Package of the EURO-BASIN programme.

Decrease in diatom palatability contributes to bloom formation in the Western English Channel

The aim of this paper is to investigate the role of phytoplankton nutritional status in the formation of the spring bloom regularly observed at the station L4 in the Western English Channel. Using a modelling approach, we tested the hypothesis that the increase in light from winter to spring induces a decrease in diatom nutritional status (i.e., an increase in the C:N and C:P ratios), thereby reducing their palatability and allowing them to bloom. To this end, a formulation describing the Stoichiometric Modulation of Predation (SMP) has been implemented in a simplified version of the European Regional Seas Ecosystem Model (ERSEM). The model was coupled with the General Ocean Turbulence Model (GOTM), implemented at the station L4 and run for 10 years (2000-.2009). Simulated carbon to nutrient ratios in diatoms were analysed in relation to microzooplankton biomass, grazing and assimilation efficiency. The model reproduced in situ data evolutions and showed the importance of microzooplankton grazing in controlling the early onset of the bloom. Simulation results supported our hypothesis and provided a conceptual model explaining the formation of the diatom spring bloom in the investigated area. However, additional data describing the microzooplankton grazing impact and the variation of carbon to nutrient ratios inside phytoplanktonic cells are required to further validate the proposed mechanisms.

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.

Modelling impacts and recovery in benthic communities exposed to localised high CO2

Regulations pertaining to carbon dioxide capture with offshore storage (CCS) require an understanding of the potential localised environmental impacts and demonstrably suitable monitoring practices. This study uses a marine ecosystem model to examine a comprehensive range of hypothetical CO2 leakage scenarios, quantifying both impact and recovery time within the benthic system. Whilst significant mortalities and long recovery times were projected for the larger and longer term scenarios, shorter-term or low level exposures lead to reduced projected impacts. This suggests that efficient monitoring and leak mitigation strategies, coupled with appropriate selection of storage sites can effectively limit concerns regarding localised environmental impacts from CCS. The feedbacks and interactions between physiological and ecological responses simulated reveal that benthic responses to CO2 leakage could be complex. This type of modelling investigation can aid the understanding of impact potential, the role of benthic community recovery and inform the design of baseline and monitoring surveys.

Modelling impacts and recovery in benthic communities exposed to localised high CO2

Regulations pertaining to carbon dioxide capture with offshore storage (CCS) require an understanding of the potential localised environmental impacts and demonstrably suitable monitoring practices. This study uses a marine ecosystem model to examine a comprehensive range of hypothetical CO2 leakage scenarios, quantifying both impact and recovery time within the benthic system. Whilst significant mortalities and long recovery times were projected for the larger and longer term scenarios, shorter-term or low level exposures lead to reduced projected impacts. This suggests that efficient monitoring and leak mitigation strategies, coupled with appropriate selection of storage sites can effectively limit concerns regarding localised environmental impacts from CCS. The feedbacks and interactions between physiological and ecological responses simulated reveal that benthic responses to CO2 leakage could be complex. This type of modelling investigation can aid the understanding of impact potential, the role of benthic community recovery and inform the design of baseline and monitoring surveys.

Carbon fluxes and pelagic ecosystem dynamics near two western Antarctic Peninsula Adélie penguin colonies: an inverse model approach

An inverse food-web model for the western Antarctic Peninsula (WAP) pelagic food web was constrained with data from Palmer Long Term Ecological Research (PAL-LTER) project annual austral summer sampling cruises. Model solutions were generated for 2 regions with Adelie penguin Pygoscelis adeliae colonies presenting different population trends (a northern and a southern colony) for a 12 yr period (1995-2006). Counter to the standard paradigm, comparisons of carbon flow through bacteria, microzooplankton, and krill showed that the diatom-krill-top predator food chain is not the dominant pathway for organic carbon exchanges. The food web is more complex, including significant contributions by microzooplankton and the microbial loop. Using both inverse model results and network indices, it appears that in the northern WAP the food web is dominated by the microbial food web, with a temporal trend toward its increasing importance. The dominant pathway for the southern WAP food web varies from year to year, with no detectable temporal trend toward dominance of microzooplankton versus krill. In addition, sensitivity analyses indicated that the northern colony of Adelie penguins, whose population size has been declining over the past 35 yr, appears to have sufficient krill during summer to sustain its basic metabolic needs and rear chicks, suggesting the importance of other processes in regulating the Adelie population decline.