Looking beyond stratification: a model-based analysis of the biological drivers of oxygen deficiency in the North Sea

Low oxygen conditions, often referred to as oxy- gen deficiency, occur regularly in the North Sea, a temperate European shelf sea. Stratification represents a major process regulating the seasonal dynamics of bottom oxygen, yet, low- est oxygen conditions in the North Sea do not occur in the regions of strongest stratification. This suggests that stratifi- cation is an important prerequisite for oxygen deficiency, but that the complex interaction between hydrodynamics and the biological processes drives its evolution. In this study we use the ecosystem model HAMSOM- ECOHAM to provide a general characterisation of the dif- ferent zones of the North Sea with respect to oxygen, and to quantify the impact of the different physical and biological factors driving the oxygen dynamics inside the entire sub- thermocline volume and directly above the bottom. With respect to oxygen dynamics, the North Sea can be subdivided into three different zones: (1) a highly produc- tive, non-stratified coastal zone, (2) a productive, season- ally stratified zone with a small sub-thermocline volume, and (3) a productive, seasonally stratified zone with a large sub- thermocline volume. Type 2 reveals the highest susceptibility to oxygen deficiency due to sufficiently long stratification pe- riods (textgreater60 days) accompanied by high surface productivity resulting in high biological consumption, and a small sub- thermocline volume implying both a small initial oxygen in- ventory and a strong influence of the biological consumption on the oxygen concentration. Year-to-year variations in the oxygen conditions are caused by variations in primary production, while spatial dif- ferences can be attributed to differences in stratification and water depth. The large sub-thermocline volume dominates the oxygen dynamics in the northern central and northern North Sea and makes this region insusceptible to oxygen de- ficiency. In the southern North Sea the strong tidal mixing inhibits the development of seasonal stratification which pro- tects this area from the evolution of low oxygen conditions. In contrast, the southern central North Sea is highly suscep- tible to low oxygen conditions (type 2). We furthermore show that benthic diagenetic processes represent the main oxygen consumers in the bottom layer, consistently accounting for more than 50% of the overall consumption. Thus, primary production followed by rem- ineralisation of organic matter under stratified conditions constitutes the main driver for the evolution of oxygen defi- ciency in the southern central North Sea. By providing these valuable insights, we show that ecosystem models can be a useful tool for the interpretation of observations and the es- timation of the impact of anthropogenic drivers on the North Sea oxygen conditions

On Using Lagrangian Drift Simulations to Aid Interpretation of in situ Monitoring Data

The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fmars.2021.666653/full#supplementary-materia

A novel modeling approach to quantify the influence of nitrogen inputs on the oxygen dynamics of the North Sea

Oxygen (O2) deficiency, i.e., dissolved O2 concentrations below 6 mg O2 L-1, is a common feature in the southern North Sea. Its evolution is governed mainly by the presence of seasonal stratification and production of organic matter, which is subsequently degraded under O2 consumption. The latter is strongly influenced by riverine nutrient loads, i.e., nitrogen (N) and phosphorus (P). As riverine P loads have been reduced significantly over the past decades, this study aims for the quantification of the influence of riverine and non-riverine N inputs on the O2 dynamics in the southern North Sea. For this purpose, we present an approach to expand a nutrient-tagging technique for physical-biogeochemical models - often referred to as 'trans-boundary nutrient transports' (TBNT) - by introducing a direct link to the O2 dynamics. We apply the expanded TBNT to the physical-biogeochemical model system HAMSOM-ECOHAM and focus our analysis on N-related O2 consumption in the southern North Sea during 2000-2014. The analysis reveals that near-bottom O2 consumption in the southern North Sea is strongly influenced by the N supply from the North Atlantic across the northern shelf edge. However, riverine N sources - especially the Dutch, German and British rivers - as well as the atmosphere also play an important role. In the region with lowest simulated O2 concentrations (around 56 °N, 6.5 °E), riverine N on average contributes 39% to overall near-bottom O2 consumption during seasonal stratification. Here, the German and the large Dutch rivers constitute the highest riverine contributions (11% and 10%, respectively). At a site in the Oyster Grounds (around 54.5 °N, 4 °E), the average riverine contribution adds up to 41%, even exceeding that of the North Atlantic. Here, highest riverine contributions can be attributed to the Dutch and British rivers adding up to almost 28% on average. The atmospheric contribution results in 13%. Our results emphasize the importance of anthropogenic N inputs and seasonal stratification for the O2 conditions in the southern North Sea. They further suggest that reductions in the riverine and atmospheric N inputs may have a relevant positive effect on the O2 levels in this region

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

Variations in the elemental ratio of organic matter in the central Baltic Sea: Part II – Sensitivities of annual mass flux estimates to model parameter variations

Highlights: • Sensitivities of annual carbon (C), nitrogen (N) and phosphorus (P) flux estimates to parameter variations are determined. • Model parameters that specify annual inventories are compared with those that determine timing and magnitude of bloom events. • Seven model parameters are of primary importance, affecting C, N and P budgets simultaneously. • Nine parameters have negligible effects on annual budget estimates and on seasonal trajectories. • Parameter categorization provides important prior information for parameter optimization in the central Baltic Sea. Abstract: This study describes a sensitivity analysis that allows the parameters of a one-dimensional ecosystem model to be ranked according to their specificity in determining biochemical key fluxes. Key fluxes of interest are annual (a) total production (TP), (b) remineralization above the halocline (RM), and (c) export at 50 m (EX) at the Baltic Sea monitoring site BY15 located in the Gotland Deep basin. The model resolves mass flux of carbon (C), nitrogen (N), and phosphorous (P), while considering nitrogen fixation explicitly. Our first null hypothesis is that the variation of the value of every single model parameter affects each annual C, N, and P budget simultaneously. Our second null hypothesis states that the variation of every parameter value induces changes at least in either of the annual C, N or P budgets. Our analyses falsify both null hypotheses and reveal that 8 out of 36 parameters must be regarded redundant, as their variation neither alter annual key fluxes nor produce considerable time-shifts in model trajectories at the respective site. Seven parameters were found to induce substantial changes in annual C, N, and P flux estimates simultaneously. The assimilation efficiency of zooplankton turned out to be of vital importance. This parameter discriminates between the assimilation and destruction of algal prey during grazing. The fraction of unassimilated dead algal cells is critical for the amount of organic matter exported out of the euphotic zone. The maximum cellular N:C quota of diazotrophs and the degradation/hydrolysis rate of detrital carbon are two parameters that will likely remain unconstrained by time series data, but both affect the annual C budget considerably. Overall, our detailed specification of model sensitivities to parameter variations will facilitate the formulation of a well-posed inverse problem for the estimation of C, N and P fluxes from stock observations at the Gotland Deep

A Novel Modeling Approach to Quantify the Influence of Nitrogen Inputs on the Oxygen Dynamics of the North Sea

Oxygen (O2) deficiency, i.e., dissolved O2 concentrations below 6 mg O2 L−1, is a common feature in the southern North Sea. Its evolution is governed mainly by the presence of seasonal stratification and production of organic matter, which is subsequently degraded under O2 consumption. The latter is strongly influenced by riverine nutrient loads, i.e., nitrogen (N) and phosphorus (P). As riverine P loads have been reduced significantly over the past decades, this study aims for the quantification of the influence of riverine and non-riverine N inputs on the O2 dynamics in the southern North Sea. For this purpose, we present an approach to expand a nutrient-tagging technique for physical-biogeochemical models — often referred to as ‘trans-boundary nutrient transports’ (TBNT) — by introducing a direct link to the O2 dynamics. We apply the expanded TBNT to the physical-biogeochemical model system HAMSOM-ECOHAM and focus our analysis on N-related O2 consumption in the southern North Sea during 2000–2014. The analysis reveals that near-bottom O2 consumption in the southern North Sea is strongly influenced by the N supply from the North Atlantic across the northern shelf edge. However, riverine N sources — especially the Dutch, German and British rivers — as well as the atmosphere also play an important role. In the region with lowest simulated O2 concentrations (around 56 °N, 6.5 °E), riverine N on average contributes 39% to overall near-bottom O2 consumption during seasonal stratification. Here, the German and the large Dutch rivers constitute the highest riverine contributions (11% and 10%, respectively). At a site in the Oyster Grounds (around 54.5 °N, 4 °E), the average riverine contribution adds up to 41%, even exceeding that of the North Atlantic. Here, highest riverine contributions can be attributed to the Dutch and British rivers adding up to almost 28% on average. The atmospheric contribution results in 13%. Our results emphasize the importance of anthropogenic N inputs and seasonal stratification for the O2 conditions in the southern North Sea. They further suggest that reductions in the riverine and atmospheric N inputs may have a relevant positive effect on the O2 levels in this region

Variations in the elemental ratio of organic matter in the central Baltic Sea: Part I—Linking primary production to remineralization

Highlights: • Elemental C:N:P variations of organic matter are simulated at monitoring site BY15. • No N2 fixation needed to explain observed PO4PO4 and pCO2pCO2 levels after spring bloom. • Model features relevance of DOP production and remineralization for N2 fixation. • Model estimates of annual N2 fixation are View the MathML source297±24mmolNm-2a-1. • Model estimates of annual total production are View the MathML source14.16±0.71molCm-2a-1. Abstract: For most marine ecosystems the growth of diazotrophic cyanobacteria and the associated amount of nitrogen fixation are regulated by the availability of phosphorus. The intensity of summer blooms of nitrogen (N2) fixing algae in the Baltic Sea is assumed to be determinable from a surplus of dissolved inorganic phosphorus (DIP) that remains after the spring bloom has ended. But this surplus DIP concentration is observed to continuously decrease at times when no appreciable nitrogen fixation is measured. This peculiarity is currently discussed and has afforded different model interpretations for the Baltic Sea. In our study we propose a dynamical model solution that explains these observations with variations of the elemental carbon-to-nitrogen-to-phosphorus (C:N:P) ratio during distinct periods of organic matter production and remineralization. The biogeochemical model resolves seasonal C, N and P fluxes with depth at the Baltic Sea monitoring site BY15, based on three assumptions: (1) DIP is utilized by algae though not needed for immediate growth, (2) the uptake of dissolved inorganic nitrogen (DIN) is hampered when the algae׳s phosphorus (P) quota is low, and (3) carbon assimilation continues at times of nutrient depletion. Model results describe observed temporal variations of DIN, DIP and chlorophyll-a concentrations along with partial pressure of carbon dioxide (pCO2)(pCO2). In contrast to other model studies, our solution does not require N2 fixation to occur shortly after the spring bloom to explain DIP drawdown and pCO2pCO2 levels. Model estimates of annual N2 fixation are View the MathML source297±24mmolNm-2a-1. Estimates of total production are View the MathML source14200±700mmolCm-2a-1, View the MathML source1400±70mmolNm-2a-1, and View the MathML source114±5mmolPm-2a-1 for the upper 50 m. The models C, N and P fluxes disclose preferential remineralization of P and of organic N that was introduced via N2 fixation. Our results are in support of the idea that P uptake by phytoplankton during the spring bloom contributes to the consecutive availability of labile dissolved organic phosphorus (LDOP). The LDOP is retained within upper layers and its remineralization affects algal growth in summer, during periods of noticeable N2 fixation

Parameterisation of a zooplankton population model for Pseudocalanus elongatus using stage durations from laboratory experiments

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