Search

Although the terms 'health' and 'healthy' are often applied to marine ecosystems and communicate information about holistic condition (e.g. as required by the Ecosystem Approach), their meaning is unclear. Ecosystems have been understood in various ways, from non-interacting populations of species to complex integrated systems. Health has been seen as a metaphor, an indicator that aggregates over system components, or a non-localized emergent system property. After a review, we define good ecosystem health as: 'the condition of a system that is self-maintaining, vigorous, resilient to externally imposed pressures, and able to sustain services to humans. It contains healthy organisms and populations, and adequate functional diversity and functional response diversity. All expected trophic levels are present and well interconnected, and there is good spatial connectivity amongst subsystems.' We equate this condition with good ecological or environmental status, e.g. as referred to by recent EU Directives. Resilience is central to health, but difficult to measure directly. Ecosystems under anthropogenic pressure are at risk of losing resilience, and thus of suffering regime shifts and loss of services. For monitoring whole ecosystems, we propose an approach based on 'trajectories in ecosystem state space', illustrated with time-series from the northwestern North Sea. Change is visualized as Euclidian distance from an arbitrary reference state. Variability about a trend in distance is used as a proxy for inverse resilience. We identify the need for institutional support for long time-series to underpin this approach, and for research to establish state space co-ordinates for systems in good health. Changes in the northern North Sea, 1958-2008, plotted in a state space defined by the breeding success of kittiwakes, abundance of copepods Calanus spp., and simulated annual primary production.© Inter-Research 2013. www.int-res.com.

The abundance and productivity of phytoplankton and bacterioplankton in the region of Jones Bank (Celtic Sea) were assessed in relation to potential physical and chemical drivers of the marine microbial community during July 2008 on RRS James Cook cruise JC25. A major storm dominated the early part of the cruise; this influenced the microbial community with a decrease in diatom abundance being evident due to mixing of cells to sub euphotic zone depths. In the euphotic zone, phytoplankton was dominated by Phaeocystis cells which, with time, became constrained within a chlorophyll maximum that was located near the top of the nutricline. Multivariate statistical analyses found that temporal changes in diatom and dinoflagellate communities were significantly related to water temperature (diatoms) and nitrate concentration (dinoflagellates). Post storm, primary productivity was greatest in the Phaeocystis dominated 2–20 μm size fraction, but with no obvious topographically generated differences between on and off bank stations. Use of a mathematical model demonstrated the likely enhancement of productivity by the storm event, and also that this storm would have masked any topographic influence on productivity. The primary production:bacterial production ratio decreased significantly with time during the cruise, suggesting active competition by bacteria with phytoplankton for mineral nutrients. This was confirmed by a deck board enrichment experiment that demonstrated a rapid increase in bacterial productivity following supplementation of the euphotic zone microbial community with nutrient rich water collected below the mixed layer.

Microplankton plays a vital part in marine ecosystems, and its importance has been recognized by the inclusion of microplankton community composition in regulatory frameworks such as the European Water Framework Directive and the Marine Strategy Framework Directive as an indicator of ecological status. Quantitative techniques are therefore required to assess the environmental status of the microplankton in a water body. Here we demonstrate the use of a method known as the microplankton index PI(mp) to evaluate changes in the microplankton community of the west coast Scottish Sea Loch Creran. Microplankton in this fjord has been studied since the 1970s, providing a data set spanning four decades. Our analysis compares an arbitrarily chosen reference period between 1979 and 1981 with a period between 2011 and 2013 and demonstrates that between these two periods community structure has changed considerably with a substantial drop in the numbers of observed diatoms accompanied by a rise in the number of autotrophic/mixotrophic dinoflagellates as well as an increase in the potentially toxin producing genus Pseudo-nitzschia and that these are related to changes in both the intensity and timing of local patterns of precipitation. The PI(mp) is shown to be a useful and robust method to visualize and quantify changes in the underlying structure of the microplankton community and is a powerful addition to the toolbox of techniques needed to determine the health of our seas.

There is growing interest in carbon stocks and flows in seagrass ecosystems, but recent global reviews suggest a paucity of studies from Africa. This paper reviews work on seagrass productivity, biomass and sediment carbon in Africa. Most work was conducted in East Africa with a major geographical gap in West Africa. The mean above-ground, below-ground and total biomasses from all studies were 174.4, 474.6 and 514 g DW m-2, respectively with a global range of 461–738 g DW m-2. Mean annual production rate was 913 g DW m-2 year-1 (global range 816–1012 g DW m-2 year-1). No studies were found giving sediment organic carbon, demonstrating a major gap in seagrass blue carbon work. Given the small numbers of relevant papers and the large geographical areas left undescribed in Africa, any conclusions remain tentative and much remains to be done on seagrass studies in Africa.

A nutrient-manipulation mesocosm study was conducted in Norwegian coastal waters to determine the effect on a nitrogen-limited microbial food web of changes in the relative concentrations of dissolved organic and inorganic nitrogen (N). Four replicated treatments were studied: no N addition, inorganic N, organic N, or a 50 : 50 mix of inorganic and organic N. Comprehensive abundance, biomass, and rate measurements were made over the 20-d experiment. The form of N available influenced species composition, succession, and the efficiency of carbon (C) incorporation. Inorganic N generated a bloom of the diatom Leptocylindrus danicus, even though silicon concentrations were below 1 mmol L 21 . The c-proteobacteria initially competed effectively with diatoms for inorganic N, exhibiting high productivity and escaping grazing control. Organic N, when available alone, was utilized more slowly by the microbial community, with diatom growth, fueled by regenerated N, achieving a lower peak abundance. A more diverse diatom species assemblage was also evident. The c-proteobacteria were less prevalent and net bacterial productivity was initially lower when the N source was organic and, in contrast to the response following inorganic N addition, the bacterial community was initially grazer controlled. A rapid succession of heterotrophic grazers occurred but with differences in species and their contribution to biomass between treatments. The efficiency of C-biomass production was lower when the N source was purely organic. However, when both inorganic and organic N were available, biomass yield was greater than the mean of the two single N source treatments.

Despite a tendency for the complexity of physical–biological models to increase, simple coupled models remain useful for some applications and can provide insights into crucial links between physical and biological processes. This argument is illustrated with an account of a simple 3-box model intended to help assess the capacity of fjords to assimilate nutrients from fish farms. The model, a dynamic version of the UK “Comprehensive Studies Task Team” (CSTT) steady-state model for eutrophication, was applied to Loch Creran (Scottish Western Highlands) and was implemented using Stella 8 and tested using historical data from 1975 (before the installation of a salmon farm) and field data collected in 2003, during the period of operation of the farm. The model's biological state variables are chlorophyll, dissolved inorganic nitrogen (DIN) and dissolved inorganic phosphorus (DIP), and it includes a simple run-off model to convert rainfall into river discharge. The physical processes involved in exchange between the loch and the adjacent waters of the Firth of Lorne were parameterised as a constant daily exchange rate. Between 1975 and 2003, local inputs of nutrient increased but, despite this, there was little apparent increase in nutrient concentrations in the loch, and observed chlorophyll concentrations decreased substantially. Model simulations of chlorophyll and DIN agreed well with observations in 1975, as did DIN simulations in 2003. However, simulated chlorophyll was overestimated in 2003. Some of the agreement between observations and simulations come from the use of observed boundary conditions to force the model. However, even when boundary conditions are subtracted from simulations and observations, the simulations in most cases retain a significant correlation with observations, demonstrating that the model's ‘interior’ processes do add to its ability to replicate conditions in the loch.

A manipulative field experiment was conducted at Blackness in the Firth of Forth, Scotland, to study the effects of differences in microphytobenthos abundance on sediment stability and macrofauna. Opaque and transparent Perspex was used to construct shaded and controlled treatments, respectively. Shading resulted in significantly lower levels of chlorophyll a recorded from the sediment. Shaded treatments showed significant reductions in populations of Macoma balthica, Hydrobia ulvae and Corophium volutator, 3 of the dominant species at this site. Largely as a result of changes in the abundance of these species, multivariate analyses showed significant differences between shaded and control communities. The impacts of shading on individual species depended on the sediment characteristics. There were no significant differences between treatments in recorded sediment accretion/erosion rates. These results demonstrate the importance of microphytobenthos for the deposit feeders M. balthica, H. ulvae and C. volutator. The absence of any effect on sediment stability might result from the reciprocal, opposite effects of macrofauna and microphytobenthos on sediment stability cancelling each other out in this experiment.

Nitrate loading to coastal waters has increased over recent decades while silicon loading has remained relatively constant or decreased. As the N:Si ratio in coastal waters shifts due to these anthropogenic influences, silicate limitation of diatom biomass may become a feature of the biogeochemistry in coastal waters especially in regions of reduced exchange. Two sets of nutrient enrichment mesocosm experiments were conducted in successive years using a natural planktonic assemblage obtained from the Trondheimsfjord, Norway. The inorganic nutrient concentrations at the start of the experiments were manipulated to give a variety of N:Si concentrations at ratios representative of current and possible future values, should N loading continue. In June 1999 experiments were conducted with a gradient of inorganic N:Si ratios (1:2, 1:1, 2:1, 4:1) to investigate the influence of low and high N:Si ratio conditions and to determine the conditions that would generate Si limitation of diatom growth. In June 2000, based on 1999 data, highly replicated experiments were conducted at N:Si ratios of 1:1 and 4:1 which were expected to result in N and Si limitation of diatom growth, respectively; statistical differences in cellular composition were recorded. N limitation of diatom biomass increase was observed under the three lowest N:Si ratios: particulate carbon (C) accumulation continued to occur following N exhaustion resulting in an increase in the organic C:N ratio. Silicate limitation of diatom biomass increase only occurred at the highest N:Si ratio of 4:1. Silicate exhaustion was followed by continued nitrate uptake for several days, at a slower rate than previously. The resulting increase in organic N was accompanied by an increase in organic C such that the C:N ratio of the organic material at the highest N:Si ratio failed to increase to the extent observed under the N limited conditions. Statistically significant differences in chlorophyll-a yield per unit nitrate, C:chlorophyll-a ratios, C:N ratio and diatom cell yield per unit nitrate or Si were observed in Si compared to N limited conditions. All mesocosms became dominated numerically and in terms of biomass by the diatom Skeletonema costatum. The potential implications of changing N and Si regimes in coastal waters are discussed.

In order to study the effect of the nitrogen:silicon (N:Si) ratio on temperate microplankton food webs, mesocosm experiments were conducted in Trondheim (Norway) using two different ratios (molar ratios of 1:1 and 4:1). With the exception of diatoms, the increase in abundance of all microbial groups [phototrophic nanoflagellates, autotrophic dinflagellates, bacteria, heterotrophic nanoflagellates (HNAN), heterotrophic microflagellates and ciliates] was significantly greater in the high N:Si treatment. Midway through the experiment, HNAN biovolume exceeded that of bacteria in the high N:Si treatment, indicating strong top-down grazing control. Heterotrophic microflagellate biovolume exceeded ciliate biovolume under both nutrient regimes. However, heterotrophic plankton failed to respond rapidly to increased diatom biomass. The heterotrophic:autotrophic biovolume ratio remained

Regions of Restricted Exchange (RREs) are an important feature of the European coastline. They are historically preferred sites for human settlement and aquaculture and their ecosystems, and consequent human use, may be at risk from eutrophication. The OAERRE project (EVK3-CT1999-0002) concerns 'Oceanographic Applications to Eutrophication in Regions of Restricted Exchange'. It began in July 2000, and studies six sites. Four of these sites are fjords: Kongsfjorden (west coast of Spitzbergen); Gullmaren (Skagerrak coast of Sweden); Himmerfjarden (Baltic coast of Sweden); and the Firth of Clyde (west coast of Scotland). Two are bays sheltered by sand bars: Golfe de Fos (French Mediterranean); and Ria Formosa (Portuguese Algarve). Together they exemplify a range of hydrographic and enrichment conditions. The project aims to understand the physical, biogeochemical and biological processes, and their interactions, that determine the trophic status of these coastal marine RRE through the development of simple screening models to define, predict and assess eutrophication. This paper introduces the sites and describes the component parts of a basic screening model and its application to each site using historical data. The model forms the starting point for the OAERRE project and views an RRE as a well-mixed box, exchanging with the sea at a daily rate E determined by physical processes, and converting nutrient to phytoplankton chlorophyll at a fixed yield q. It thus uses nutrient levels to estimate maximum biomass; these preliminary results are discussed in relation to objective criteria used to assess trophic status. The influence of factors such as grazing and vertical mixing on key parameters in the screening model are further studied using simulations of a complex 'research' model for the Firth of Clyde. The future development of screening models in general and within OAERRE in particular is discussed. In addition, the paper looks ahead with a broad discussion of progress in the scientific understanding of eutrophication and the legal and socioeconomic issues that need to be taken into account in managing the trophic status of RREs.

Nutrient manipulation experiments were conducted on a natural planktonic community in outdoor mesocosms. Inorganic nitrogen (N) and silicon (Si) were added to achieve N[ratio ]Si ratios of 1[ratio ]1 and 4[ratio ]1. Total particulate carbon (PC) biomass of the microbial assemblage was determined by elemental analysis. Cell volume measurements by microscope on individual components of the community (bacteria, diatoms, photosynthetic nanoflagellates, heterotrophic nanoflagellates, dinoflagellates and ciliates) were also made. We applied published C[ratio ]volume relationships to determine the volume estimated C content (CBV) of these microbial groups and hence of the total assemblage. The total CBV and total PC were compared to test the applicability of C[ratio ]volume relationships under different nutrient regimes both before and after nutrient exhaustion. For initial N[ratio ]Si ratios of 1[ratio ]1, prior to nutrient exhaustion, the relationship between CBV and PC was linear with a gradient of approximately 1, (0.99±0.06), indicating that the published C[ratio ]volume relationships accurately predicted the C content of the microbial assemblage. For N[ratio ]Si ratios of 4[ratio ]1, a linear relationship was again evident between CBV and PC (slope: 1.36±0.08). However, statistical comparison using a general linear model indicated that the gradient of this relationship differed significantly from that when the N[ratio ]Si ratio was 1[ratio ]1, and hence CBV overestimated elemental C. For both N[ratio ]Si ratios, subsequent to nutrient exhaustion (N or Si), and hence when the diatom fraction of the microbial assemblage was in yield-limited post-exponential phase, the two measures of biomass were not well correlated. This indicated that measured cytoplasmic cell volume was a poor indicator of C biomass within the microbial assemblage in nutrient-deplete conditions.

Primary production was measured at a series of stations in austral summer 1996 in the vicin- ity of South Georgia. Five stations were occupied along a 700 km transect southeastwards towards the western tip of South Georgia. Three stations were located north of the Antarctic Polar Front (PF) with mean primary production of 0.41 g C m -2 d -1 compared to 0.54 g C m -2 d -1 for the stations south of the front. Three stations were in the shelf break and off-shelf region to the east of South Georgia where the lowest rates of primary production were recorded (mean 0.34 g C m -2 d -1 ). At a further 4 stations north- west of the island, primary production was significantly greater: 0.8 to 2.5 g C m -2 d -1 (mean 1.5 g C m -2 d -1 ). Photoinhibition was a marked feature of the production profiles, resulting in the underestimation of column production by 4 to 16%. An assessment is made of the primary production required to main- tain a representative assemblage of metazooplankton around South Georgia composed of 30 g fresh mass (FM) m -2 krill and 20 g FM m -2 copepods. The total C requirement of both groups at maximum growth rate is estimated as 0.93 g C m -2 d -1 , with 0.54 g C m -2 d -1 required to maintain basal metabolism (i.e. for zero growth). Local primary production rates are well in excess of the C demands for basal metabolism by the metazooplankton, and likewise the highest regional values exceed those for maxi- mum metazooplankton growth. It is also estimated that the krill biomass required to sustain higher predators during their breeding season could be supported by local production at plausible maximum krill growth rate. In particular, high primary production found in this study to the northwest of South Georgia appears to be adequate to sustain maximum growth rate by krill when their biomass is 30 g FM m -2 , when krill production will exceed predator removal. It is already well known that advection from higher latitudes is important in supplying energy to the South Georgia system. This study adds to the evidence that locally enhanced primary production is also important in supporting the local food web.

An understanding of undesirable disturbance to the balance of organisms is needed to diagnose marine eutrophication as defined by EU Directives and OSPAR. This review summarizes the findings of the UK Defra-funded Undesirable Disturbance Study Team, which concluded that 'an undesirable disturbance is a perturbation of a marine ecosystem that appreciably degrades the health or threatens the sustainable human use of that ecosystem'. A methodology is proposed for detecting disturbance of temperate salt-water communities dominated by phytoplanktonic or phytobenthic primary producers. It relies on monitoring indicators of ecosystem structure and vigour, which are components of health. Undesirable disturbance can be diagnosed by accumulating evidence of ecohydrodynamic type-specific changes in: (i) bulk indicators; (ii) frequency statistics; (iii) flux measurements; (iv) structural indicators; and (v) indicator species. These are exemplified by (i) chlorophyll, transparency, dissolved oxygen, and opportunistic seaweed cover; (ii) HABs frequency; (iii) primary production; (iv) benthic and planktonic 'trophic indices'; (v) seagrasses and Nephrops norvegicus. Ecological Quality Objectives are proposed for some of these. Linking the diagnosis to eutrophication requires correlation of changes with nutrient enrichment. The methodology, which requires the development of a plankton community index and emphasizes the importance of primary production as an indicator of vigour, can be harmonized with the EU Water Framework Directive and OSPAR's Strategy to Combat Eutrophication.

Oceanic dimethylsulfide (DMS), the main natural source of sulfur to the global atmosphere, is suggested to play a key role in the interaction between marine biota and climate. Its biochemical precursor is dimethylsulfoniopropionate (DMSP), a globally distributed, intracellular constituent in marine phytoplankton. During a multidisciplinary Lagrangian experiment in the subpolar North Atlantic, we determined the fluxes of DMSP and DMS through phytoplankton, microzooplankton, and bacterioplankton and compared them with concurrent carbon and sulfur fluxes through primary and secondary productions, grazing, and release and use of dissolved organic matter. We found that DMSP and derivatives contributed most (48-100%) of the sulfur fluxes and 5-15% of the carbon fluxes. Our findings highlight DMSP as a prominent player in pelagic biogeochemical pumps, especially as a major carrier in organic sulfur cycling. Also, our results illustrate the key role played by microzooplankton and heterotrophic bacteria (hence the microbial food web) in controlling the amount of phytoplanktonic DMSP that ultimately vents to the atmosphere in the form of DMS.

Jickells, T.D. ... et. al.-- 34 pages, 6 tables, 10 figures We report the results of an experiment in the Northeast Atlantic in which sulphur hexafluoride (SF6) was released within an eddy and the behaviour of trace gases, nutrients and productivity followed within a Lagrangian framework over a period of 24 days. Measurements were also made in the air above the eddy in order to estimate air–sea exchange rates for some components. The physical, biological and biogeochemical properties of the eddy resemble those of other eddies studied in this area, suggesting that the results we report may be applicable beyond the specific eddy studied. During a period of low wind speed at the start of the experiment, we are able to quantitatively describe and balance the nutrient and carbon budgets for the eddy. We also report concentrations of various trace gases in the region which are similar to those observed in other studies and we estimate exchange rates for several trace gases. We show that the importance of gas exchange over other loss terms varies with time and also varies for the different gases. We show that the various trace gases considered (CO2, dimethyl sulphide (DMS), N2O, CH4, non-methane-hydrocarbons, methyl bromide, methyl iodide and volatile selenium species) are all influenced by physical and biological processes, but the overall distribution and temporal variability of individual gases are different to one another. A storm disrupted the stratification in the eddy during the experiment, resulting in enhanced nutrient supply to surface waters, enhanced gas exchange rates and a change in plankton community, which we quantify, although overall productivity was little changed. Emphasis is placed on the regularity of storms in the temperate ocean and the importance of these stochastic processes in such systems This is contribution no. ACP175 of the NERC ACSOE Thematic Programme. NERC grants GST/02/1276 supported the work of Jickells and Spokes, GST/02/1278 the work of Liss and Nightingale together with core funding to the Plymouth Marine Laboratory (PML) while the ACSOE core programme supported Broadgate and the shiptime costs, NERC grant GR9/3467 to G.S. supported Moncoiffé and Gilpin. Steinke was supported by NERC grant GR3/10956. The University of Newcastle upon Tyne Research Committee provided funding to support T. Frost. The work of Simó and Pedrós-Alió was supported by the Spanish McyT grant MAR97-1885-E