Autotrophic Stoichiometry Emerging from Optimality and Variable Co-limitation

Autotrophic organisms reveal an astounding flexibility in their elemental stoichiometry, with potentially major implications on biogeochemical cycles and ecological functioning. Notwithstanding, stoichiometric regulation, and co-limitation by multiple resources in autotrophs were in the past often described by heuristic formulations. In this study, we present a mechanistic model of autotroph growth, which features two major improvements over the existing schemes. First, we introduce the concept of metabolic network independence that defines the degree of phase-locking between accessory machines. Network independence is in particular suggested to be proportional to protein synthesis capability as quantified by variable intracellular N:C. Consequently, the degree of co-limitation becomes variable, contrasting with the dichotomous debate on the use of Liebig's law or the product rule, standing for constantly low and high co-limitation, respectively. Second, we resolve dynamic protein partitioning to light harvesting, carboxylation processes, and to an arbitrary number of nutrient acquisition machineries, as well as instantaneous activity regulation of nutrient uptake. For all regulatory processes we assume growth rate optimality, here extended by an explicit consideration of indirect feed-back effects. The combination of network independence and optimal regulation displays unprecedented skill in reproducing rich stoichiometric patterns collected from a large number of published chemostat experiments. This high skill indicates (1) that the current paradigm of fixed co-limitation is a critical short-coming of conventional models, and (2) that stoichiometric flexibility in autotrophs possibly reflects an optimality strategy. Numerical experiments furthermore show that regulatory mechanisms homogenize the effect of multiple stressors. Extended optimality alleviates the effect of the most limiting resource(s) while down-regulating machineries for the less limiting ones, which induces an ubiquitous response surface of growth rate over ambient resource levels. Our approach constitutes a basis for improved mechanistic understanding and modeling of acclimative processes in autotrophic organisms. It hence may serve future experimental and theoretical investigations on the role of those processes in aquatic and terrestrial ecosystems

The large scale impact of offshore wind farm structures on pelagic primary productivity in the southern North Sea

The increasing demand for renewable energy is projected to result in a 40-fold increase in offshore wind electricity in the European Union by 2030. Despite a great number of local impact studies for selected marine populations, the regional ecosystem impacts of offshore wind farm structures are not yet well assessed nor understood. Our study investigates whether the accumulation of epifauna, dominated by the filter feeder Mytilus edulis (blue mussel), on turbine structures affects pelagic primary productivity and ecosystem functioning in the southern North Sea. We estimate the anthropogenically increased potential distribution based on the current projections of turbine locations and reported patterns of M. edulis settlement. This distribution is integrated through the Modular Coupling System for Shelves and Coasts to state-of-the-art hydrodynamic and ecosystem models. Our simulations reveal non-negligible potential changes in regional annual primary productivity of up to 8% within the offshore wind farm area, and induced maximal increases of the same magnitude in daily productivity also far from the wind farms. Our setup and modular coupling are effective tools for system scale studies of other environmental changes arising from large-scale offshore wind-farming such as ocean physics and distributions of pelagic top predators.Comment: 17 pages, 6 figures, re-revised manuscript submitted to Hydrobiologi

Modular System for Shelves and Coasts (MOSSCO v1.0) - a flexible and multi-component framework for coupled coastal ocean ecosystem modelling

Shelf and coastal sea processes extend from the atmosphere through the water column and into the sea bed. These processes are driven by physical, chemical, and biological interactions at local scales, and they are influenced by transport and cross strong spatial gradients. The linkages between domains and many different processes are not adequately described in current model systems. Their limited integration level in part reflects lacking modularity and flexibility; this shortcoming hinders the exchange of data and model components and has historically imposed supremacy of specific physical driver models. We here present the Modular System for Shelves and Coasts (MOSSCO, http://www.mossco.de), a novel domain and process coupling system tailored---but not limited--- to the coupling challenges of and applications in the coastal ocean. MOSSCO builds on the existing coupling technology Earth System Modeling Framework and on the Framework for Aquatic Biogeochemical Models, thereby creating a unique level of modularity in both domain and process coupling; the new framework adds rich metadata, flexible scheduling, configurations that allow several tens of models to be coupled, and tested setups for coastal coupled applications. That way, MOSSCO addresses the technology needs of a growing marine coastal Earth System community that encompasses very different disciplines, numerical tools, and research questions.Comment: 30 pages, 6 figures, submitted to Geoscientific Model Development Discussion

The Predominant Processes Controlling Vertical Nutrient and Suspended Matter Fluxes across Domains - Using the New MOSSCO System from Coastal Sea Sediments up to the Atmosphere

Integrated Modeling of Hydro-System

Shape matters: the relationship between cell geometry and diversity in phytoplankton

Size and shape profoundly influence an organism’s ecophysiological performance and evolutionary fitness, suggesting a link between morphology and diversity. However, not much is known about how body shape is related to taxonomic richness, especially in microbes. Here we analyse global datasets of unicellular marine phytoplankton, a major group of primary producers with an exceptional diversity of cell sizes and shapes and, additionally, heterotrophic protists. Using two measures of cell shape elongation, we quantify taxonomic diversity as a function of cell size and shape. We find that cells of intermediate volume have the greatest shape variation, from oblate to extremely elongated forms, while small and large cells are mostly compact (e.g. spherical or cubic). Taxonomic diversity is strongly related to cell elongation and cell volume, together explaining up to 92% of total variance. Taxonomic diversity decays exponentially with cell elongation and displays a log-normal dependence on cell volume, peaking for intermediate-volume cells with compact shapes. These previously unreported broad patterns in phytoplankton diversity reveal selective pressures and ecophysiological constraints on the geometry of phytoplankton cells which may improve our understanding of marine ecology and the evolutionary rules of life

Temporal declines in Wadden Sea phytoplankton cell volumes observed within and across species

Cell size is a master trait in the functional ecology of phytoplankton correlating with numerous morphological, physiological, and life-cycle characteristics of species that constrain their nutrient use, growth, and edibility. In contrast to well-known spatial patterns in cell size at macroecological scales or temporal changes in experimental contexts, few data sets allow testing temporal changes in cell sizes within ecosystems. To analyze the temporal changes of intraspecific and community-wide cell size, we use the phytoplankton data derived from the Lower Saxony Wadden Sea monitoring program, which comprises sample- and species-specific measurements of cell volume from 1710 samples collected over 14 yr. We find significant reductions in both the cell volume of most species and the weighted mean cell size of communities. Mainly diatoms showed this decline, whereas the size of dinoflagellates seemed to be less responsive. The magnitude of the trend indicates that cell volumes are about 30% smaller now than a decade ago. This interannual trend is overlayed by seasonal cycles with smaller cells typically observed in summer. In the subset of samples including environmental conditions, small community cell size was strongly related to high temperatures and low total phosphorus concentration. We conclude that cell size captures ongoing changes in phytoplankton communities beyond the changes in species composition. In addition, based on the changes in species biovolumes revealed by our analysis, we warn that using standard cell size values in phytoplankton assessment will not only miss temporal changes in size, but also lead to systematic errors in biomass estimates over time

A model-based projection of historical state of a coastal ecosystem : relevance of phytoplankton stoichiometry

We employed a coupled physical-biogeochemical modelling framework for the reconstruction of the historic (H), pre-industrial state of a coastal system, the German Bight (southeastern North Sea), and we investigated its differences with the recent, control (C) state of the system. According to our findings: i) average winter concentrations of dissolved inorganic nitrogen and phosphorus (DIN and DIP) concentrations at the surface are ∼70–90% and ∼50–70% lower in the H state than in the C state within the nearshore waters, and differences gradually diminish towards off-shore waters; ii) differences in average growing season chlorophyll a (Chl) concentrations at the surface between the two states are mostly less than 50%; iii) in the off-shore areas, Chl concentrations in the deeper layers are affected less than in the surface layers; iv) reductions in phytoplankton carbon (C) biomass under the H state are weaker than those in Chl, due to the generally lower Chl:C ratios; v) in some areas the differences in growth rates between the two states are negligible, due to the compensation by lower light limitation under the H state, which in turn explains the lower Chl:C ratios; vi) zooplankton biomass, and hence the grazing pressure on phytoplankton is lower under the H state. This trophic decoupling is caused by the low nutritional quality (i.e., low N:C and P:C) of phytoplankton. These results call for increased attention to the relevance of the acclimation capacity and stoichiometric flexibility of phytoplankton for the prediction of their response to environmental change