From plankton to top predators: bottom-up control of a marine food web across four trophic levels

1. Abundant mid-trophic pelagic fish often play a central role in marine ecosystems, both as links between zooplankton and top predators and as important fishery targets. In the North Sea, the lesser sandeel occupies this position, being the main prey of many bird, mammal and fish predators and the target of a major industrial fishery. However, since 2003, sandeel landings have decreased by > 50%, and many sandeel-dependent seabirds experienced breeding failures in 2004. 2. Despite the major economic implications, current understanding of the regulation of key constituents of this ecosystem is poor. Sandeel abundance may be regulated 'bottom-up' by food abundance, often thought to be under climatic control, or 'top-down' by natural or fishery predation. We tested predictions from these two hypotheses by combining unique long-term data sets (1973–2003) on seabird breeding productivity from the Isle of May, SE Scotland, and plankton and fish larvae from the Continuous Plankton Recorder survey. We also tested whether seabird breeding productivity was more tightly linked to sandeel biomass or quality (size) of individual fish. 3. The biomass of larval sandeels increased two- to threefold over the study period and was positively associated with proxies of the abundance of their plankton prey. Breeding productivity of four seabirds bringing multiple prey items to their offspring was positively related to sandeel larval biomass with a 1-year lag, indicating dependence on 1-year-old fish, but in one species bringing individual fish it was strongly associated with the size of adult sandeels. 4. These links are consistent with bottom-up ecosystem regulation and, with evidence from previous studies, indicate how climate-driven changes in plankton communities can affect top predators and potentially human fisheries through the dynamics of key mid-trophic fish. However, the failing recruitment to adult sandeel stocks and the exceptionally low seabird breeding productivity in 2004 were not associated with low sandeel larval biomass in 2003, so other mechanisms (e.g. predation, lack of suitable food after metamorphosis) must have been important in this case. Understanding ecosystem regulation is extremely important for predicting the fate of keystone species, such as sandeels, and their predators.

Biological impacts of enhanced alkalinity in Carcinus maenas

Further steps are needed to establish feasible alleviation strategies that are able to reduce the impacts of ocean acidification, whilst ensuring minimal biological side-effects in the process. Whilst there is a growing body of literature on the biological impacts of many other carbon dioxide reduction techniques, seemingly little is known about enhanced alkalinity. For this reason, we investigated the potential physiological impacts of using chemical sequestration as an alleviation strategy. In a controlled experiment, Carcinus maenas were acutely exposed to concentrations of Ca(OH)2 that would be required to reverse the decline in ocean surface pH and return it to pre-industrial levels. Acute exposure significantly affected all individuals' acid-base balance resulting in slight respiratory alkalosis and hyperkalemia, which was strongest in mature females. Although the trigger for both of these responses is currently unclear, this study has shown that alkalinity addition does alter acid-base balance in this comparatively robust crustacean species.

Use Of Micro-Stereology And Quantitative Cyto-Chemistry To Determine The Effects Of Crude Oil-Derived Aromatic-Hydrocarbons On Lysosomal Structure And Function In A Marine Bivalve Mollusk Mytilus-Edulis

The marine bivalve mollusc,Mytilus edulis (blue mussel), is a noted accumulator of many environmental pollutants and is increasingly used for the chemical and biological assessment of environmental impact. The toxic effects of crude oil-derived aromatic hydrocarbons (30 μg/l total hydrocarbons) on the lysosomal-vacuolar system of the digestive cells have been investigated in cryostat sections of hexane-frozen digestive glands. Exposure to aromatic hydrocarbons reduced the cytochemically determined latency of lysosomal β-N-acetylhexosaminidase; lysosomal volume density and surface density increased while the numerical density decreased. Experimental exposure resulted in the formation of very large lysosomes which are believed to be largely autophagic in function and these results indicate a significant structural and functional disturbance of digestive cell lysosomes in response to hydrocarbons.

Potential consequences of climate change for primary production and fish production in large marine ecosystems

Existing methods to predict the effects of climate change on the biomass and production of marine communities are predicated on modelling the interactions and dynamics of individual species, a very challenging approach when interactions and distributions are changing and little is known about the ecological mechanisms driving the responses of many species. An informative parallel approach is to develop size-based methods. These capture the properties of food webs that describe energy flux and production at a particular size, independent of species' ecology. We couple a physical-biogeochemical model with a dynamic, size-based food web model to predict the future effects of climate change on fish biomass and production in 11 large regional shelf seas, with and without fishing effects. Changes in potential fish production are shown to most strongly mirror changes in phytoplankton production. We project declines of 30-60% in potential fish production across some important areas of tropical shelf and upwelling seas, most notably in the eastern Indo-Pacific, the northern Humboldt and the North Canary Current. Conversely, in some areas of the high latitude shelf seas, the production of pelagic predators was projected to increase by 28-89%.

Potential Climate Change Effects on the Habitat of Antarctic Krill in the Weddell Quadrant of the Southern Ocean

Antarctic krill is a cold water species, an increasingly important fishery resource and a major prey item for many fish, birds and mammals in the Southern Ocean. The fishery and the summer foraging sites of many of these predators are concentrated between 0 degrees and 90 degrees W. Parts of this quadrant have experienced recent localised sea surface warming of up to 0.2 degrees C per decade, and projections suggest that further widespread warming of 0.27 degrees to 1.08 degrees C will occur by the late 21st century. We assessed the potential influence of this projected warming on Antarctic krill habitat with a statistical model that links growth to temperature and chlorophyll concentration. The results divide the quadrant into two zones: a band around the Antarctic Circumpolar Current in which habitat quality is particularly vulnerable to warming, and a southern area which is relatively insensitive. Our analysis suggests that the direct effects of warming could reduce the area of growth habitat by up to 20%. The reduction in growth habitat within the range of predators, such as Antarctic fur seals, that forage from breeding sites on South Georgia could be up to 55%, and the habitat's ability to support Antarctic krill biomass production within this range could be reduced by up to 68%. Sensitivity analysis suggests that the effects of a 50% change in summer chlorophyll concentration could be more significant than the direct effects of warming. A reduction in primary production could lead to further habitat degradation but, even if chlorophyll increased by 50%, projected warming would still cause some degradation of the habitat accessible to predators. While there is considerable uncertainty in these projections, they suggest that future climate change could have a significant negative effect on Antarctic krill growth habitat and, consequently, on Southern Ocean biodiversity and ecosystem services.

Transmission electron microscopy of marine crustacean eggs

Ultrastructural investigations of eggs can be important in helping to understand embryonic development. There are few transmission electron microscope studies of marine arthropod eggs, however, as they have proved difficult to fix and infiltrate with resin. Here, we describe a modification of a standard method that allows the preparation of the quite different eggs of the marine copepod, Acartia tonsa and the lobster, Homarus gammarus, for transmission electron microscopy. By using double fixation and an extended resin infiltration time we obtained good preparations for electron microscopy. We anticipate that these modifications to the standard protocol will be widely applicable and useful for the study of the eggs and early developmental stages of many marine arthropod taxa. Les recherches sur l'ultrastructure des oeufs peuvent être importantes en aidant à comprendre le développement embryonnaire. Il existe cependant peu d'études en microscopie électronique à transmission sur les oeufs d'arthropodes marins, car il est difficile de les fixer et d'y infiltrer de la résine. Dans ce travail, nous décrivons une modification de la méthode standard, qui permet la préparation pour la microscopie électronique à transmission d'oeufs aussi différents que ceux du copépode marin Acartia tonsa et du homard Homarus gammarus. En utilisant une double fixation et un temps plus long d'infiltration de la résine, nous avons obtenu de bonnes préparations pour la microscopie électronique. Nous prévoyons que ces modifications du protocole standard seront largement applicables et utiles pour l'étude des oeufs et des premiers stades de développement de nombreux taxons d'arthropodes marins.

Climate-induced effects on the meroplankton and the benthic-pelagic ecology of the North Sea

Analyses of long-term time series of North Sea plankton and sea surface temperature (SST) data reveal that the annual planktonic larval abundance of three benthic phyla, Echinodermata, Arthropoda, and Mollusca, responds positively and immediately to SST. Long-term outcomes for the planktonic abundance of these three phyla are different, however. The planktonic larvae of echinoderms and decapod crustaceans have increased in abundance from 1958 to 2005, and especially since the mid-1980s, as North Sea SST has increased. In contrast, the abundance of bivalve mollusc larvae has declined, despite the positive year-to-year relationship between temperature and bivalve larval abundance continuing to hold. We argue that the changes in meroplankton abundance, coincident with increased phytoplankton and declining holoplankton, reflect the synchronous effect of rising SST and related changes in the pelagic community on the reproduction and recruitment of many benthic marine invertebrates. Under this scenario, the long-term decline in bivalve mollusc larvae will reflect increased predation on the settled larvae and adults by benthic decapods. These alterations in the zooplankton may therefore describe an ecosystem-wide restructuring of North Sea trophic interactions.

Synergistic effects of climate and fishing in a marine ecosystem

Current climate change and overfishing are affecting the productivity and structure of marine ecosystems. This situation is unprecedented for the marine biosphere and it is essential to understand the mechanisms and pathways by which ecosystems respond. We report that climate change and overfishing are likely to be responsible for a rapid restructuring of a highly productive marine ecosystem with effects throughout the pelagos and the benthos. In the mid-1980s, climate change, consequent modifications in the North Sea plankton, and fishing, all reduced North Sea cod recruitment. In this region, production of many benthic species respond positively and immediately to temperature. Analysis of a long-term, spatially extensive biological (plankton and cod) and physical (sea surface temperature) dataset suggests that synchronous changes in cod numbers and sea temperature have established an extensive trophic cascade favoring lower trophic level groups over economic fisheries. A proliferation of jellyfish that we detect may signal the climax of these changes. This modified North Sea ecology may provide a clear indication of the synergistic consequences of coincident climate change and overfishing. The extent of the ecosystem restructuring that has occurred in the North Sea suggests we are unlikely to reverse current climate and human-induced effects through ecosystem resource management in the short term. Rather, we should understand and adapt to new ecological regimes. This implies that fisheries management policies will have to be fully integrated with the ecological consequences of climate change to prevent a similar collapse in an exploited marine ecosystem elsewhere.

Recovery or decline of the Northwestern Black Sea: A societal choice revealed by socio-ecological modelling

During recent decades anthropogenic activities have dramatically impacted the Black Sea ecosystem. High levels of riverine nutrient input during the 1970s and 1980s caused eutrophic conditions including intense algal blooms resulting in hypoxia and the subsequent collapse of benthic habitats on the northwestern shelf. Intense fishing pressure also depleted stocks of many apex predators, contributing to an increase in planktivorous fish that are now the focus of fishing efforts. Additionally, the Black Sea's ecosystem changed even further with the introduction of exotic species. Economic collapse of the surrounding socialist republics in the early 1990s resulted in decreased nutrient loading which has allowed the Black Sea ecosystem to start to recover, but under rapidly changing economic and political conditions, future recovery is uncertain. In this study we use a multidisciplinary approach to integrate information from socio-economic and ecological systems to model the effects of future development scenarios on the marine environment of the northwestern Black Sea shelf. The Driver–Pressure–State-Impact-Response framework was used to construct conceptual models, explicitly mapping impacts of socio-economic Drivers on the marine ecosystem. Bayesian belief networks (BBNs), a stochastic modelling technique, were used to quantify these causal relationships, operationalise models and assess the effects of alternative development paths on the Black Sea ecosystem. BBNs use probabilistic dependencies as a common metric, allowing the integration of quantitative and qualitative information. Under the Baseline Scenario, recovery of the Black Sea appears tenuous as the exploitation of environmental resources (agriculture, fishing and shipping) increases with continued economic development of post-Soviet countries. This results in the loss of wetlands through drainage and reclamation. Water transparency decreases as phytoplankton bloom and this deterioration in water quality leads to the degradation of coastal plant communities (Cystoseira, seagrass) and also Phyllophora habitat on the shelf. Decomposition of benthic plants results in hypoxia killing flora and fauna associated with these habitats. Ecological pressure from these factors along with constant levels of fishing activity results in target stocks remaining depleted. Of the four Alternative Scenarios, two show improvements on the Baseline ecosystem condition, with improved waste water treatment and reduced fishing pressure, while the other two show a worsening, due to increased natural resource exploitation leading to rapid reversal of any recent ecosystem recovery. From this we conclude that variations in economic policy have significant consequences for the health of the Black Sea, and ecosystem recovery is directly linked to social–economic choices.

Effects on zooplankton of a warmer ocean: Recent evidence from the Northeast Pacific

The consequences for pelagic communities of warming trends in mid and high latitude ocean regions could be substantial, but their magnitude and trajectory are not yet known. Environmental changes predicted by climate models (and beginning to be confirmed by observations) include warming and freshening of the upper ocean and reduction in the extent and duration of ice cover. One way to evaluate response scenarios is by comparing how "similar" zooplankton communities have differed among years and/or locations with differing temperature. The subarctic Pacific is a strong candidate for such comparisons, because the same mix of zooplankton species dominates over a wide range of temperature climatologies, and observations have spanned substantial temperature variability at interannual-to-decadal time scales. In this paper, we review and extend copepod abundance and phenology time series from net tow and Continuous Plankton Recorder surveys in the subarctic Northeast Pacific. The two strongest responses we have observed are latitudinal shifts in centers of abundance of many species (poleward under warm conditions), and changes in the life cycle timing of Neocalanus plumchrus in both oceanic and coastal regions (earlier by several weeks in warm years and at warmer locations). These zooplankton data, plus indices of higher trophic level responses such as reproduction, growth and survival of pelagic fish and seabirds, are all moderately-to-strongly intercorrelated (vertical bar r vertical bar = 0.25-0.8) with indices of local and basin-scale temperature anomalies. A principal components analysis of the normalized anomaly time series from 1979 to 2004 shows that a single "warm-and-low-productivity" vs. "cool-and-high-productivity" component axis accounts for over half of the variance/covariance. Prior to 1990, the scores for this component were negative ("cool" and "productive") or near zero except positive in the El Nino years 1983 and 1987. The scores were strongly and increasingly positive ("warm" and "low productivity") from 1992 to 1998; negative from 1999 to 2002; and again increasingly positive from 2003-present. We suggest that, in strongly seasonal environments, anomalously high temperature may provide misleading environmental cues that contribute to timing mismatch between life history events and the more-nearly-fixed seasonality of insolation, stratification, and food supply. Crown Copyright (c) 2007 Published by Elsevier Ltd. All rights reserved.

Trophic level asynchrony in rates of phenological change for marine, freshwater and terrestrial environments

Recent changes in the seasonal timing (phenology) of familiar biological events have been one of the most conspicuous signs of climate change. However, the lack of a standardized approach to analysing change has hampered assessment of consistency in such changes among different taxa and trophic levels and across freshwater, terrestrial and marine environments. We present a standardized assessment of 25 532 rates of phenological change for 726 UK terrestrial, freshwater and marine taxa. The majority of spring and summer events have advanced, and more rapidly than previously documented. Such consistency is indicative of shared large scale drivers. Furthermore, average rates of change have accelerated in a way that is consistent with observed warming trends. Less coherent patterns in some groups of organisms point to the agency of more local scale processes and multiple drivers. For the first time we show a broad scale signal of differential phenological change among trophic levels; across environments advances in timing were slowest for secondary consumers, thus heightening the potential risk of temporal mismatch in key trophic interactions. If current patterns and rates of phenological change are indicative of future trends, future climate warming may exacerbate trophic mismatching, further disrupting the functioning, persistence and resilience of many ecosystems and having a major impact on ecosystem services.

Improving the performance of a Mediterranean demersal fishery towards societal objectives beyond MSY

Mediterranean demersal fisheries are highly multispecific and many of their target stocks are overexploited. In addition, rocketing fuel costs and low market prices of traditionally high-value species are challenging the viability of fisheries. Here, based on the numeric results of a simulation model, we conclude that this situation can be remedied by reducing both fishing mortality and fishing costs. According to our model results, fishing effort reductions of 48–71% would improve the health of fish stocks while increasing the economic profits of Mallorca islands bottom trawl fishery to as much as 1.9 M€ (146% higher than current profits). If all fish stocks were exploited at their MSY (or below) level, the reduction in fishing effort would have to be of 71% from current values. If equilibrium profits from the fishery were to be maximized (MEY), fishing effort would need to be reduced by 48%. These results must be taken with caution due the many sources of uncertainty of our analysis. The modeling tools used to estimate these values are conditional to the adequate treatment of two sources of uncertainty that are particularly problematic in Mediterranean fisheries: insufficiently known recruitment variability and lack of periodic evaluations of the state of many species. Our results show that fishing effort reductions would produce economic yield gains after a period of transition. Further studies on the benefits of changing the size-selection pattern of fisheries, on better estimation of stock–recruitment relationships and on better quantifications of the contribution of secondary species to these fisheries, are expected to improve the scientific recommendations for Mediterranean demersal fisheries toward sustainability principles.

Estimating the economic loss of recent North Atlantic fisheries management

It is accepted that world’s fisheries are not generally exploited at their biological or their economic optimum. Most fisheries assessments focus on the biological capacity of fish stocks to respond to harvesting and few have attempted to estimate the economic efficiency at which ecosystems are exploited. The latter is important as fisheries contribute considerably to the economic development of many coastal communities. Here we estimate the overall potential economic rent for the fishing industry in the North Atlantic to be B€ 12.85, compared to current estimated profits of B€ 0.63. The difference between the potential and the net profits obtained from North Atlantic fisheries is therefore B€ 12.22. In order to increase the profits of North Atlantic fisheries to a maximum, total fish biomass would have to be rebuilt to 108 Mt (2.4 times more than present) by reducing current total fishing effort by 53%. Stochastic simulations were undertaken to estimate the uncertainty associated with the aggregate bioeconomic model that we use and we estimate the economic loss NA fisheries in a range of 2.5 and 32 billion of euro. We provide economic justification for maintaining or restoring fish stocks to above their MSY biomass levels. Our conclusions are consistent with similar global scale studies.

Implications of streamlining theory for microbial ecology

Whether a small cell, a small genome or a minimal set of chemical reactions with self-replicating properties, simplicity is beguiling. As Leonardo da Vinci reportedly said, 'simplicity is the ultimate sophistication'. Two diverging views of simplicity have emerged in accounts of symbiotic and commensal bacteria and cosmopolitan free-living bacteria with small genomes. The small genomes of obligate insect endosymbionts have been attributed to genetic drift caused by small effective population sizes (Ne). In contrast, streamlining theory attributes small cells and genomes to selection for efficient use of nutrients in populations where Ne is large and nutrients limit growth. Regardless of the cause of genome reduction, lost coding potential eventually dictates loss of function. Consequences of reductive evolution in streamlined organisms include atypical patterns of prototrophy and the absence of common regulatory systems, which have been linked to difficulty in culturing these cells. Recent evidence from metagenomics suggests that streamlining is commonplace, may broadly explain the phenomenon of the uncultured microbial majority, and might also explain the highly interdependent (connected) behavior of many microbial ecosystems. Streamlining theory is belied by the observation that many successful bacteria are large cells with complex genomes. To fully appreciate streamlining, we must look to the life histories and adaptive strategies of cells, which impose minimum requirements for complexity that vary with niche.

Basin-scale phenology and effects of climate variability on global timing of initial seaward migration of Atlantic salmon (Salmo salar)

Migrations between different habitats are key events in the lives of many organisms. Such movements involve annually recurring travel over long distances usually triggered by seasonal changes in the environment. Often, the migration is associated with travel to or from reproduction areas to regions of growth. Young anadromous Atlantic salmon (Salmo salar) emigrate from freshwater nursery areas during spring and early summer to feed and grow in the North Atlantic Ocean. The transition from the freshwater (parr') stage to the migratory stage where they descend streams and enter salt water (smolt') is characterized by morphological, physiological and behavioural changes where the timing of this parr-smolt transition is cued by photoperiod and water temperature. Environmental conditions in the freshwater habitat control the downstream migration and contribute to within- and among-river variation in migratory timing. Moreover, the timing of the freshwater emigration has likely evolved to meet environmental conditions in the ocean as these affect growth and survival of the post-smolts. Using generalized additive mixed-effects modelling, we analysed spatio-temporal variations in the dates of downstream smolt migration in 67 rivers throughout the North Atlantic during the last five decades and found that migrations were earlier in populations in the east than the west. After accounting for this spatial effect, the initiation of the downstream migration among rivers was positively associated with freshwater temperatures, up to about 10 degrees C and levelling off at higher values, and with sea-surface temperatures. Earlier migration occurred when river discharge levels were low but increasing. On average, the initiation of the smolt seaward migration has occurred 2.5days earlier per decade throughout the basin of the North Atlantic. This shift in phenology matches changes in air, river, and ocean temperatures, suggesting that Atlantic salmon emigration is responding to the current global climate changes.