Trophic network model of the Northern Adriatic Sea: Analysis of an exploited and eutrophic ecosystem

A quantitative model of the trophic network of Northern Adriatic Sea marine ecosystem during the 1990s has been constructed, with the goal of analysing its trophic structure, identifying the key trophic groups and assessing the anthropogenic impacts on the ecosystem using the Ecopath modelling protocol. The Northern Adriatic Sea is an eutrophic, shallow basin, and one of the most heavily fished areas in the Mediterranean Sea. The network aggregation into discrete trophic levels sensu Lindeman shows that low trophic levels dominate biomass and energy flows, with 40% of the total system throughput flowing out from trophic level 2. Instead, upper trophic levels appear bottom-up controlled, highly depleted and not exerting any control on the trophic network, as shown by mixed trophic impact-based analyses. Microbial loop is comparable to grazing with respect to the magnitude of flows involved, as 66% of the trophic network flows originate from detritus, which is mainly consumed by bacteria. Key trophic groups are plankton groups, macro-crustaceans and detritus, and other r-selected organisms like squids and small pelagics, which have a great influence on the ecosystem. In particular, zooplankton acts as a bottleneck for energy flows, limiting the energy from the low trophic levels effectively reaching the upper food web. The high pelagic production caused by eutrophication sustains high fishery landings and impressive discard quantities, as well as the benthic compartment. Overall, the ecosystem appears quite productive and in a stressed and developmental status. Model results and comparisons with few existing historical data suggest that the low maturity and stressed state of the Northern Adriatic Sea are not only due to natural characteristics, but mainly to anthropogenic pressures.     

海洋浮游生态系统动力学模式的研究

分析了海洋浮游生态系统动力学模式的结构,讨论了其研究现状。９０年代以来海洋浮游生态系统动力学模式的特点是：状态变量的个数较以前有所增加,从２－４个增加至３－７个生物过程的描述更加细致;注意刻画物理过程的作用,特别是垂直方向上湍流的交换作用;利用或改进原来的模式,研究新问题;模式仍以一维为主,同时三维耦合模式也得取发展和应用;区域仍以北大西洋为主,也出现了研究赤道太平洋、阿拉伯海、东海等海区的工作。     

Predicting the distribution of deep-sea vulnerable marine ecosystems using high-resolution data: Considerations and novel approaches

Little is known about species distribution patterns in deep-sea environments, primarily because sampling surveys in the high seas are expensive and time consuming. The increasing need to manage and protect vulnerable marine ecosystems, such as cold-water corals, has motivated the use of predictive modelling tools, which produce continuous maps of potential species or habitat distribution from limited point observations and full coverage environmental data. Rapid advances in acoustic remote sensing, oceanographic modelling and sampling technology now provide high quality datasets, facilitating model development with high spatial detail. This paper provides a short overview of existing methodologies for predicting deep-sea benthic species distribution, and illustrates emerging issues related to spatial and thematic data resolution, and the use of transect-derived species distribution data. In order to enhance the ecological relevance and reliability of deep-sea species distribution models, novel techniques are presented based on a case study predicting the distribution of the cold-water coral Lophelia pertusa in three carbonate mound provinces in Irish waters. Specifically, the study evaluates (1) the capacity of newly developed high-resolution (250 m grid cell size) hydrodynamic variables to explain local scale cold-water coral distribution patterns, (2) the potential value of species occurrence proportion data to maintain semi-quantitative information of coral prevalence (i.e. coverage) and sampling effort per grid cell within the response variable, and (3) mixed effect modelling to deal with spatially grouped transect data. The study shows that predictive models using vertical and horizontal flow parameters perform significantly better than models based on terrain parameters only. Semi-quantitative proportion data may decrease model uncertainty and increase model reliability, and provide a fruitful avenue of research for analysing large quantities of video data in a detailed yet time-efficient manner. The study concludes with an outlook of how species distribution models could improve our understanding of vulnerable marine ecosystem functioning and processes in the deep sea.     

Abrupt transitions of the top-down controlled Black Sea pelagic ecosystem during 1960–2000: Evidence for regime-shifts under strong fishery exploitation and nutrient enrichment modulated by climate-induced variations

Functioning of the Black Sea ecosystem has profoundly changed since the early 1970s under cumulative effects of excessive nutrient enrichment, strong cooling/warming, over-exploitation of pelagic fish stocks, and population outbreak of gelatinous carnivores. Applying a set of criteria to the long-term (1960–2000) ecological time-series data, the present study demonstrates that the Black Sea ecosystem was reorganised during this transition phase in different forms of top-down controlled food web structure through successive regime-shifts of distinct ecological properties. The Secchi disc depth, oxic–anoxic interface zone, dissolved oxygen and hydrogen sulphide concentrations also exhibit abrupt transition between their alternate regimes, and indicate tight coupling between the lower trophic food web structure and the biogeochemical pump in terms of regime-shift events.The first shift, in 1973–1974, marks a switch from large predatory fish to small planktivore fish-controlled system, which persisted until 1989 in the form of increasing small pelagic and phytoplankton biomass and decreasing zooplankton biomass. The increase in phytoplankton biomass is further supported by a bottom-up contribution due to the cumulative response to high anthropogenic nutrient load and the concurrent shift of the physical system to the “cold climate regime” following its ∼20-year persistence in the “warm climate regime”. The end of the 1980s signifies the depletion of small planktivores and the transition to a gelatinous carnivore-controlled system. By the end of the 1990s, small planktivore populations take over control of the system again. Concomitantly, their top-down pressure when combined with diminishing anthropogenic nutrient load and more limited nutrient supply into the surface waters due to stabilizing effects of relatively warm winter conditions switched the “high production” regime of phytoplankton to its background “low production” regime.The Black Sea regime-shifts appear to be sporadic events forced by strong transient decadal perturbations, and therefore differ from the multi-decadal scale cyclical events observed in pelagic ocean ecosystems under low-frequency climatic forcing. The Black Sea observations illustrate that eutrophication and extreme fishery exploitation can indeed induce hysteresis in large marine ecosystems, when they can exert sufficiently strong forcing onto the system. They further illustrate the link between the disruption of the top predators, proliferation of new predator stocks, and regime-shift events. Examples of these features have been reported for some aquatic ecosystems, but are extremely limited for large marine ecosystems.     

How well do ecosystem indicators communicate the effects of anthropogenic eutrophication?

Anthropogenic eutrophication affects the Mediterranean, Black, North and Baltic Seas to various extents. Responses to nutrient loading and methods of monitoring relevant indicators vary regionally, hindering interpretation of ecosystem state changes and preventing a straightforward pan-European assessment of eutrophication symptoms. Here we summarize responses to nutrient enrichment in Europe's seas, comparing existing time-series of selected pelagic (phytoplankton biomass and community composition, turbidity, N:P ratio) and benthic (macro flora and faunal communities, bottom oxygen condition) indicators based on their effectiveness in assessing eutrophication effects. Our results suggest that the Black Sea and Northern Adriatic appear to be recovering from eutrophication due to economic reorganization in the Black Sea catchment and nutrient abatement measures in the case of the Northern Adriatic. The Baltic is most strongly impacted by eutrophication due to its limited exchange and the prevalence of nutrient recycling. Eutrophication in the North Sea is primarily a coastal problem, but may be exacerbated by climatic changes. Indicator interpretation is strongly dependent on sea-specific knowledge of ecosystem characteristics, and no single indicator can be employed to adequately compare eutrophication state between European seas. Communicating eutrophication-related information to policy-makers could be facilitated through the use of consistent indicator selection and monitoring methodologies across European seas. This work is discussed in the context of the European Commission's recently published Marine Strategy Directive.     

An integrated approach for assessing the relative significance of human pressures and environmental forcing on the status of Large Marine Ecosystems

An ecosystem approach to the management of the marine environment has received considerable attention over recent years. However, there are few examples which demonstrate its practical implementation. Much of this relates to the history of existing marine monitoring and assessment programmes which (for many countries) are sectoral, making it difficult to integrate monitoring data and knowledge across programmes at the operational level.To address this, a scientific expert group, under the auspices of the International Council for the Exploration of the Sea (ICES), prepared a plan for how ICES could contribute to the development of an Integrated Ecosystem Assessment (IEA) for the North Sea by undertaking a pilot study utilising marine monitoring data. This paper presents the main findings arising from the expert group and in particular it sets out one possible integrated approach for assessing the relative significance of environmental forcing and fishing pressure on the ecological status of the North Sea, it then compares the findings with assessments made of other Large Marine Ecosystems (LMEs).We define the North Sea ecosystem on the basis of 114 state and pressure variables resolved as annual averages between 1983 and 2003 and at the spatial scale of ICES rectangles. The paper presents results of integrated time-series and spatial analysis which identifies and explains significant spatial and temporal gradients in the data. For example, a significant shift in the status of the North Sea ecosystem (based upon 114 state-pressure variables) is identified to have occurred around 1993. This corresponds to previously documented shifts in the environmental conditions (particularly sea surface temperature) and changes in the distribution of key species of plankton (Calanus sp.), both reported to have occurred in 1989. The difference in specific timing between reported regime shifts for the North Sea may be explained, in part, by time-lag dependencies in the trophic structure of the ecosystem with shifts in higher trophic levels occurring later than 1989.By examining the connection (or relatedness) between ecosystem components (e.g. environment, plankton, fish, fishery and seabirds) for the identified regime states (1983–1993; 1993–2003) we conclude that both the North Sea pelagic and benthic parts of the ecosystem were predominantly top-down (fishery) controlled between 1983 and 1993, whereas between 1993 and 2003 the pelagic stocks shifted to a state responding mainly to bottom-up (environment) influences. However, for the demersal fish stocks between 1993 and 2003 top-down (fishery) pressure dominated even though over this period significant reductions in fishing pressure occurred. The present analysis, therefore, provides further evidence in support of the need for precautionary management measures taken in relation to setting fishery quotas.     

A model study on carbon cycle and phytoplankton dynamical processes in the Bohai Sea

The carbon cycle of lower trophic level in the Bohai Sea is studied with a three-dimensional biological and physical coupled model. The influences of the processes (including horizontal advection, river nutrient load, active transport etc. ) on the phytoplankton biomass and its evolution are estimated. The Bohai Sea is a weak sink of the CO2 in the atmosphere. During the cycle, 13.7% of the gross production of the phytoplankton enter the higher trophic level and 76.8 % of it are consumed by the respiration itself. The nutrient reproduction comes mainly from the internal biogeochemical loop and the rem-ineralization is an important mechanism of the nutrient transfer from organic form to inorganic. Horizontal advection decreases the total biomass and the eutrophication in some sea areas. Change in the nutrient load of a river can only adjust the local system near its estuary. Controlling the input of the nutrient, which limits the alga growth, can be very useful in lessening the phytoplankton biomass.     

Towards end-to-end models for investigating the effects of climate and fishing in marine ecosystems

End-to-end models that represent ecosystem components from primary producers to top predators, linked through trophic interactions and affected by the abiotic environment, are expected to provide valuable tools for assessing the effects of climate change and fishing on ecosystem dynamics. Here, we review the main process-based approaches used for marine ecosystem modelling, focusing on the extent of the food web modelled, the forcing factors considered, the trophic processes represented, as well as the potential use and further development of the models. We consider models of a subset of the food web, models which represent the first attempts to couple low and high trophic levels, integrated models of the whole ecosystem, and size spectrum models. Comparisons within and among these groups of models highlight the preferential use of functional groups at low trophic levels and species at higher trophic levels and the different ways in which the models account for abiotic processes. The model comparisons also highlight the importance of choosing an appropriate spatial dimension for representing organism dynamics. Many of the reviewed models could be extended by adding components and by ensuring that the full life cycles of species components are represented, but end-to-end models should provide full coverage of ecosystem components, the integration of physical and biological processes at different scales and two-way interactions between ecosystem components. We suggest that this is best achieved by coupling models, but there are very few existing cases where the coupling supports true two-way interaction. The advantages of coupling models are that the extent of discretization and representation can be targeted to the part of the food web being considered, making their development time- and cost-effective. Processes such as predation can be coupled to allow the propagation of forcing factors effects up and down the food web. However, there needs to be a stronger focus on enabling two-way interaction, carefully selecting the key functional groups and species, reconciling different time and space scales and the methods of converting between energy, nutrients and mass.     

Ecosystem structure and resilience—A comparison between the Norwegian and the Barents Sea

Abundance and biomass of the most important fish species inhabited the Barents and Norwegian Sea ecosystems have shown considerable fluctuations over the last decades. These fluctuations connected with fishing pressure resulted in the trophic structure alterations of the ecosystems. Resilience and other theoretical concepts (top-down, wasp-waste and bottom-up control, trophic cascades) were viewed to examine different response of the Norwegian and Barents Sea ecosystems on disturbing forces. Differences in the trophic structure and functioning of Barents and Norwegian Sea ecosystems as well as factors that might influence the resilience of the marine ecosystems, including climatic fluctuation, variations in prey and predator species abundance, alterations in their regular migrations, and fishing exploitation were also considered. The trophic chain lengths in the deep Norwegian Sea are shorter, and energy transfer occurs mainly through the pelagic fish/invertebrates communities. The shallow Barents Sea is characterized by longer trophic chains, providing more energy flow into their benthic assemblages. The trophic mechanisms observed in the Norwegian Sea food webs dominated by the top-down control, i.e. the past removal of Norwegian Spring spawning followed by zooplankton development and intrusion of blue whiting and mackerel into the area. The wasp-waist response is shown to be the most pronounced effect in the Barents Sea, related to the position of capelin in the ecosystem; large fluctuations in the capelin abundance have been strengthened by intensive fishery. Closer links between ecological and fisheries sciences are needed to elaborate and test various food webs and multispecies models available.     

Quantitative model of trophic interactions in Beibu Gulf ecosystem in the northern South China Sea

1IntroductionThe Beibu Gulf is a natural semiclosed conti-nental sea of the South China Sea,which is situatedat17°00′~21°45′N,105°40′~110°10′E,and sur-rounded by China and Vietnam(see Fig.1).It hasa subtropic monsoon climate with an average winter     

Does ecological redundancy maintain functioning of marine benthos on centennial to millennial time scales?

Predicting the ability of the biosphere to continue to deliver ecosystem services in the face of biodiversity loss and environmental change is a major challenge. The results of short‐term and small‐scale experimental studies are both equivocal and difficult to extrapolate from. In this study we use data on benthic palaeocommunities covering 4,000,000 years (in the Late Jurassic when temperate coastal seas in NW Europe experienced fluctuations in oxygenation). The biological traits associated with each species in the palaeocommunities were combined to index the delivery of ecological functions. Five ecosystem functions were examined: food for large mobile predators, biogenic habitat provision, nutrient recycling/regeneration, inorganic carbon sequestration and food‐web dynamics. In modern systems these ecological functions underpin ecosystem services that are important for human well‐being. Our results show that the supply of food for higher predators was remarkably constant during the 4,000,000 years, suggesting that redundancy amongst species in the assemblage drives the biodiversity–ecosystem function (BEF) relationship. By contrast, the provision of biogenic habitat varied with the occurrence of a relatively few taxa, a pattern consistent with a rivet type model of BEF. For nutrient regeneration, carbon sequestration and food‐web dynamics the patterns were complex and suggestive of an idiosyncratic model of BEF. To our knowledge this is the first study to quantify ecological functioning through deep time and demonstrates the utility of this approach to understanding long‐term patterns of BEF in both ancient and contemporary marine ecosystems. The delivery of all five ecological functions studied became increasingly variable as the regional climate became drier, thus modifying the supply of terrigenous nutrient inputs.     

Separate and interactive effects of eutrophication and climate variables on the ecosystem elements of the Gulf of Riga

There is currently a critical knowledge gap in how eutrophication and climate variables separately and interactively impact the dynamics of marine ecosystems. Based on long-term monitoring data we quantified the separate and combined impacts of nutrient loading, temperature, salinity, and wind conditions on zooplankton, zoobenthos and fish inhabiting a brackish water ecosystem in the Gulf of Riga. Changes in zoobenthos communities and herring stock were largely explained by climate variables. Zooplankton species were related to both eutrophication and climate variables, and models combining all environmental variables explained additional variation in zooplankton data compared to the separate models of climate and eutrophication. This suggests that zoobenthos communities and herring stock are largely driven by weather conditions, whereas the combined effect of weather and nutrient loads are likely the cause for dynamic zooplankton communities in the Gulf of Riga.     

Considering ecosystem-based fisheries management in the California Current

Recognizing that all management decisions have impacts on the ecosystem being exploited, an ecosystem-based approach to management seeks to better inform these decisions with knowledge of ecosystem structure, processes and functions. For marine fisheries in the California Current, along the West Coast of North America, such an approach must take into greater consideration the constantly changing climate-driven physical and biological interactions in the ecosystem, the trophic relationships between fished and unfished elements of the food web, the adaptation potential of life history diversity, and the role of humans as both predators and competitors. This paper reviews fisheries-based ecosystem tools, insights, and management concepts, and presents a transitional means of implementing an ecosystem-based approach to managing US fisheries in the California Current based on current scientific knowledge and interpretation of existing law.     

Ecological performance indicators in the North Sea: development and application

Different signals of distress from the North Sea ecosystems started a discussion on the protection of the ecosystem at the third North Sea Ministers Conference in 1990. This was followed by a number of workshops on ecological indicators under the auspices of the North Sea Task Force and the Oslo and Paris Commissions (OSPAR). In 1997 the member countries around the North Sea agreed to develop and apply an ecosystem's approach in the management programs for both the North Sea fisheries and the marine environment. Following this agreement the identification of the ecological qualities objectives for the North Sea ecosystem started. Further work in this direction has lead to the setting up of various national projects geared towards creating a better understanding of the North Sea ecosystem.In the Netherlands, two departments (Water management and Nature and fisheries management) from the Ministry of Transport, Public works and Water Management (V&W) and the Ministry of Agriculture, Nature and Fisheries Management (LNV) agreed to integrate their policies for the management of the Netherlands section of the North Sea. The results from these projects (started in 1996) should enable the policy makers and managers to formulate policies that could lead to the creation of a balance between the effects of human activities and the preservation of the natural qualities of the North Sea ecosystem.One of the projects is strictly designed to develop ecological indicators for the Dutch part of the North Sea. These indicators are based on various monitoring data and related in one way or another with human activities. In this way it is possible to evaluate the effects of human impacts on the ecosystem. Long-term changes in the ecological performance of the ecosystem can be elucidated by the use of the ecological indicators. Thereby making it possible for the policy makers and the managers of the NCP to evaluate the effects of their policies and management recommendations on the quality of the ecosystem.This paper describes the process and the steps taken by the authorities in the countries around the North Sea to develop a set of ecological indicators for the management of the North Sea ecosystem. Focus will be made on the efforts made in the Netherlands to develop indicators that did provide the basis for the development of Water and Nature conservation policies and new management strategies for maintaining the sustainable use of the Dutch section of the North Sea.     

Long-term simulation of the eutrophication of the North Sea: temporal development of nutrients, chlorophyll and primary production in comparison to observations

The ecosystem model ERSEM II has been used to hindcast the development of the ecosystem of the North Sea during the years 1955 to 1993. The simulation was driven by the box-aggregated output from a general circulation model of the North Sea of corresponding duration; radiation, river inputs, atmospheric input and boundary conditions at the borders to the Atlantic Ocean and to the Baltic Sea were applied as realistically as possible. The general features of the eutrophication process are reproduced in the hindcast: the coastal areas show strong changes in nutrient concentrations in the hindcast as well as in the observations. Eutrophication not only shows up in the nutrient concentrations, but also in primary production. The simulated spatial distributions of phosphate, nitrate and primary production compare well with the observed ones. In addition, the hindcast simulates considerable trend-like changes of the nutrients in the southern part of the North Sea, where the nutrients are transported from the continental coastal strip to the southern central North Sea. The line from the river Humber to southern Norway separates the region of noticeable anthropogenic influence of riverine and atmospheric input from the northern area, which is mainly influenced by the Atlantic nutrient inflow. The observed annual cycles in the central and northern North Sea are quite well reproduced by the hindcast. The comparison of the hindcast with the long-term observations at two sites in the continental coastal zone of the North Sea shows that the long-term behaviour of phosphate, nitrate and silicate is simulated well. Primary production is increased in summers during the main period of eutrophication, 1975 to 1989, in the hindcast and in the observations. The flagellates at Helgoland, however, experience much more pronounced annual cycles with much less interannual variability in the hindcast than in the observations.     

Trophic interactions in the coastal ecosystem of Sri Lanka: An ECOPATH preliminary approach

This study attempts to assemble and summarize existing information in order to build a general representation of the trophic interactions within the shallow coastal ecosystem of Sri Lanka. A multispecific ecosystem-based approach on trophic relationships and their possible variations was performed using ECOPATH. Thirty-nine functional groups were considered representing all trophic levels in the food web.     

热带典型珊瑚岛礁海洋牧场花刺参底播增殖容量及其生态效应预测

为改善热带珊瑚岛礁型海洋牧场的珊瑚礁生境,实现生物资源的养护和渔业资源的产出功能,在对海参等高值经济种开展底播增殖前,科学评估其生态容量是防止引发海洋牧场生态风险的重要保证。运用生态系统模型法评估了三亚蜈支洲岛热带珊瑚岛礁海洋牧场花刺参(Stichopus monotuberculatus)的底播增殖容量。根据2020~2021年蜈支洲岛海洋牧场近岛区渔业资源调查与环境因子数据,运用Ecopath with Ecosim 6.6软件构建了该海域的生态系统营养通道模型。研究表明:生态系统各功能组营养级范围介于1~3.52,系统的食物网结构以牧食食物链为主,总能流中有43%的能量来源于碎屑功能组,其在系统总能流中有重要地位。系统的总平均能量传递效率为9.353%,略低于林德曼能量传递效率(10%)。总初级生产量/总呼吸量为3.726,总初级生产量/总生物量为28.834,系统连接指数为0.256,杂食性指数为0.120,系统Finn''s循环指数和平均路径长度分别为2.485%和2.379,表明近岛区生态系统食物网结构较为简单,且系统稳定性和成熟度偏低,易受外界干扰。根据模型评估的花刺参增殖生态容量为110.21 t/km²,是现存量的206 倍,有较大增殖空间,并且达到生态容量后碎屑组的能量再循环利用效率将显著增加,营养级结构能得到进一步优化,系统稳定性及成熟度将有所提高。基于研究结果,可适当采捕与花刺参生态位相近的生物,同时增殖放流其他处于不同营养层次的经济种,从而减少种间竞争,有效利用系统冗余能量,进而扩大花刺参的生态容量,实现海洋牧场的健康可持续发展。     

A review of the NEMURO and NEMURO.FISH models and their application to marine ecosystem investigations

The evolution of the North Pacific Ecosystem Model for Understanding Regional Oceanography (NEMURO) family of models to study marine ecosystems is reviewed. Applications throughout the North Pacific have shown the models to be robust and to be able to reproduce 1D, 2D and 3D components of nutrient, carbon cycle and biogeochemical cycles as well as aspects of the lower trophic levels ecosystem (phyto- and zooplankton). NEMURO For Including Saury and Herring, an extension that includes higher trophic levels, can be run uncoupled or coupled to NEMURO. In the uncoupled mode, the growth and weight of an individual fish is computed using plankton densities simulated by NEMURO but with no feedback between fish consumption and plankton mortality. In the coupled mode, the feeding, growth and weight of a representative fish are computed, and prey removals due to feeding by fish appear as mortality terms on the prey. The NEMURO family of models continues to evolve, including effects of the microbial loop and iron limitation at lower trophic levels, and full life cycle, multi-species and multi-generational simulations at higher trophic levels. We outline perspectives for future end-to-end modeling efforts that can be used to study marine ecosystems in response to global environmental change.     

Climatological annual cycles of nutrients and chlorophyll in the North Sea

A large amount of nutrient and chlorophyll data from the North Sea were compiled and organised in a research data base to produce annual cycles on a relatively fine spatial resolution of 1° in each horizontal direction. The data originate from many different sources and were partly provided by the ECOMOD data base of the Institut fur Meereskunde in Hamburg and partly by ICES in Copenhagen to cover the time range from 1950 to 1994. While the annual cycles of nutrients and chlorophyll derived for the continental coastal zone are representative for the decade 1984–1993 only, those for the remaining parts of the North Sea may be considered climatological annual cycles based on data from more than four decades. The composite data set of climatological annual cycles of medians and their climatological ranges is well suited to serve for validational and forcing purposes for ecosystem models of the North Sea, which have a resolution larger than or equal to 1° in both longitude and latitude. The annual cycles of the macronutrients and chlorophyll presented here for 1° × 1° squares in the North Sea show especially that sufficient observational data exist to provide initial, forcing and validational data for the simulations with the 130-box setup (ND130) of the ecosystem model ERSEM. The annual cycles presented give a clear picture for the whole of the North Sea. The highest concentrations occur at the continental coasts as a result of continued river input, which is added to the ongoing atmospheric input over the North Sea. Also, from the Atlantic Ocean water with relatively high nutrient concentrations enters the North Sea via the northern boundary. In the productive areas on and around the Dogger Bank nutrient concentrations are lower than in the other parts of the North Sea, even in winter. The areas with seasonal stratification have very different annual cycles in the upper (0–30 m) and lower layers (30 m-bottom). The shallow boxes are fully mixed and exhibit a relatively fast increase of nutrient concentrations caused by summer regeneration of nutrients.