ALBERT

Description: We examined the RNA and DNA concentration of field-caught scallops Chlamys islandica, maintained in suspended cultures at 15 and 30 m depth, and scallops from a wild population at 50 to 60 m in Kobbefjord, southwest Greenland. General relations between RNA and DNA concentrations and individual shell height were established, and we found that the RNA:DNA ratio (RD) worked well as a standardisation of the RNA concentration independent of size and sex. During an experimental period of 14 mo, we observed a pronounced seasonal pattern in RD and mass growth, and differences between depths. Even though the period with high levels of RD reflected the growth season relatively well, RD was a poor predictor of individual mass growth rates of C. islandica. However, we found a non-linear response in RD to increased food concentrations resulting in RD being up- and down-regulated at the beginning and end of the productive summer season, respectively. These results indicate that short-term dynamics in the actual mass growth rate might be controlled through regulation of ribosome activity rather than ribosome number (RNA concentration). This adaption would allow scallops to up-regulate protein synthesis more rapidly, thereby ensuring efficient utilisation of the intense peaks in food availability in coastal areas in the Arctic. Therefore, we suggest that RD in C. islandica reflects the growth potential rather than the actual growth rate. Still, the amount of unexplained variance in RD is considerable and not independent over time, suggesting the existence of unresolved mechanisms or relationships.

Description: The effect of ambient solar ultraviolet radiation (UVR) on a shallow-water (4 cm) tropical fouling community was assessed during the succession of macrobenthic species on artificial substrates at the Wong Shek fish farm, Hong Kong. The early successional communities developing under 3 radiation treatments (PAR + UV-A + UV-B = 280 to 700 nm; PAR + UV-A = 320 to 700 nm, and PAR = 400 to 700 nm) were monitored for 14 wk. A total of 8 species of algae and 8 species of invertebrates colonised the experimental tiles. During the first 8 wk of the experiments, there were no differences among treatments in diversity, percentage of cover of species and the biomass of the colonisers. During the following 6 wk, the communities exposed to UVR had lower species richness than the communities exposed to only PAR had. The species diversity (after 79 and 98 d) of the 3 treatments varied, but the total percentage of species cover and the entire community biomass were not significantly different across the experiment. Juveniles of the polychaete Hydroides elegans and the barnacle Balanus amphitrite, juveniles of the clams Perna viridis and Modiolus comptus, and the algae Enteromorpha sp., Ectocarpus sp. and Cladophora sp. were responsible for the dissimilarity between communities developed under different UVR treatments. The algae constituted a higher percentage of the cover under the full sunlight spectrum, whereas the polychaete, the barnacle and the clams were dominant in the no-UVR treatment. Our outdoor experiment revealed that UVR inhibited the settlement and decreased post-settlement survival of H. elegans. We concluded that UVR affects the composition of early successional, shallow water biofouling communities in tropical waters as well as the settlement and mortality of single species.

Description: We investigated connections between subtropical Atlantic climate variability, atmospheric conditions in the European Alpine region (45 to 47° N and 5 to 8° E) and the interannual variability of the thermal conditions in the largest body of freshwater in Western Europe (Lake Geneva). The long-term water temperature was related to climate variability by means of a multivariate regression model. Results revealed atmospheric connections that have been elusive so far, and showed that over the period from 1959 to 2000, summer thermal conditions in Lake Geneva appear tightly linked to the long-term variability of the subtropical Atlantic climate. The multivariate model revealed high skills and tight correlations, which suggest the possibility of assessing future thermal changes in Lake Geneva from the Atlantic climate variability. The implications of such climatic forcing on the functioning of the pelagic ecosystem in Lake Geneva were illustrated by analysing the long-term changes in abundance of the summer-dominant carnivorous cladocerans Bythotrephes longimanus and Leptodora kindtii during the period 1974 to 2000. Again, the multivariate model revealed high skills and excellent correlations between the interannual changes in abundance of these species and the variability of summer climate. Our approach provides a general understanding of the interrelations between large- and regional-scale climates, local environmental conditions and the ecological responses in Lake Geneva during summer, and is therefore applicable to other retrospective studies.

Description: Throughout the last 2 decades a shift from a cod- to a sprat-dominated system occurred in the upper trophic levels of the Central Baltic Sea. This was caused by a decline in the cod stock, due to recruitment failure and high fishing intensity, resulting in a decrease in predation pressure on sprat. Concurrently with the lowest cod stock size on record, sprat reached biomass values of above 2 × 106 t in 1992, being relatively stable afterwards. Besides predation mortality through cod and in recent years also an increasing fishing pressure, cannibalism on eggs may be a compensatory process limiting the reproductive success of sprat and hence contributing to the population regulation in the Central Baltic. Based on sprat stomach sampling on 21 cruises between March 1988 and July 1996 cannibalism on sprat eggs was investigated in the Bornholm Basin, one of the main spawning areas of Central Baltic sprat. Using a model of gastric evacuation to estimate daily food intake rates and a Virtual Population/Extended Survivor Analysis for computing predator population sizes, egg cannibalism rates were estimated. These were compared to egg abundance data from ichthyoplankton surveys and to preliminary estimates of seasonal egg productions. The study revealed significant interannual differences in the intensity of sprat egg cannibalism with considerable predation in 1990 to 1992 (〉15% of the egg abundance during peak spawning and 〉60% of the seasonal production) and a reduction in most recent years (〈16% of the corresponding abundance and production). As a possible reason for these differences a combination of changes in the vertical overlap of predator/prey and variability in the food environment were identified. Shortcomings of the applied methods and the possible impact of cannibalism on the reproductive success and population development of sprat in the Central Baltic Sea are discussed.

Description: It is widely assumed that the production of secondary metabolites against grazing and fouling is costly for seaweeds in terms of metabolic energy and should therefore be reduced under conditions of resource limitation. Here we tested the hypothesis that anti-herbivore defenses and bioactivity against mussels in 4 brown seaweeds from northern-central Chile will be reduced when light is limited. In a 2 wk experiment, seaweeds were kept under different low-light conditions (~76 to 99% reduction of ambient sunlight) and grazing situations. Subsequently, we tested their anti-herbivore defense against a common amphipod grazer in feeding assays with living algal tissue and reconstituted food pellets. A standard test employing the production of byssus threads by mussels was furthermore used as an indicator for deterrents in crude algal extracts. All investigated seaweeds showed decreased growth under the stepwise light reduction. Lessonia nigrescens exhibited reduced defense ability under severe low-light conditions when living tissue was offered to the amphipod, probably caused by changes in the tissue structure or in nutritional traits. In Dictyota kunthii, L. trabeculata and Macrocystis integrifolia this effect was absent. None of the investigated seaweeds showed a clear effect of light reduction on chemically mediated defenses against the mesograzer and there was no effect of light limitation on the bioactivity against mussels. Thus, against general assumptions, chemical defense in the investigated seaweeds does not appear to be reduced under severe resource limitation. Results suggest that seaweeds may use different strategies of energy allocation to cope with low-light conditions.

Description: The abundance and diversity of microorganisms on the surface of the tropical green macroalga Caulerpa racemosa var. turbinata and the effect of algal surface and waterborne compounds on fouling organisms were investigated both in laboratory and field experiments. As shown via electron microscopic enumeration, the abundance of epibiotic bacteria and diatoms on algal frond surfaces was not significantly different from the reference biofilms harvested from stones in the C. racemosa habitat. The analysis of Terminal Restriction Fragment Length Polymorphism of DNA from algal surface-associated bacterial communities revealed that despite a similar abundance of these bacteria, the community profile on algal frond surfaces differed significantly from that of inanimate, undefended substrates. These results suggest that the alga regulate the occurrence of certain bacterial ribotypes. This result was in accordance with the fact that different bacterial communities formed on the artificial substrata (i.e. Petri dishes) placed in the C. racemosa habitat and alga-free control sites. Neither C. racemosa conditioned seawater (CCW) nor hexane surface extracts affected the growth of bacterial isolates from biofilms. However, only CCW exhibited a toxic effect on the larvae of the fouling polychaete Hydroides elegans, and evoked abnormal larval development in a concentration-dependent fashion. At sublethal concentrations, the 〈1 kD fraction of CCW inhibited the larval settlement of H. elegans and the bryozoan Bugula neritina. Caulerpenyne, the prominent bioactive metabolite in the genus Caulerpa, was not detected in CCW by chromatographic procedures. Our data suggest that waterborne compounds other than caulerpenyne are involved in the chemical defense of the alga C. racemosa.

Description: The foraging modes of calanoid copepods differ in that stationary suspension-feeding is more easily detected by prey with strong escape responses (ciliates) than is ‘cruising’ or ‘ambushing’ feeding. Thus, the ability of a copepod to include heterotrophic prey in its diet may be associated with its foraging mode and, further, with its nitrogen stable isotope signature (δ15N). This is because a more carnivorous diet may be expected to result in a higher δ15N. We tested this hypothesis in a mesocosm study using a density gradient (0 to 80 ind. l-1) of calanoid copepods. We expected copepod δ15N to generally increase with decreasing copepod density because of increased food availability, and predicted stronger increases for cruising than for stationary suspension-feeding species. As an assemblage, copepods had a pronounced impact on the food web: diatoms and ciliates decreased, whereas nanoflagellates increased with increasing copepod density. As expected, Centropages hamatus, a cruising species, showed the strongest isotopic increase and also highest population growth at low copepod density, suggesting that it was the most efficient species in capturing ciliates. Temora longicornis, a stationary suspension-feeder, showed a uniform isotopic increase in all mesocosms, which we believe resulted from nutritional stress arising from poor feeding on both ciliates (too fast for ingestion by T. longicornis) and nanoflagellates (too small). However, Pseudocalanus elongatus, a species equally categorised as a stationary suspension-feeder, showed increases in its δ15N similar to those for C. hamatus. While this may indicate potential switching in its foraging mode, alternative explanations cannot be ruled out, partly because qualitative and quantitative aspects of trophic enrichment in our experiment could not be clearly separated. This study shows that consumer δ15N are difficult to interpret, even if potential food sources and aspects of the species’ biology are known, and thus emphasises the necessity for further laboratory studies to help better interpret zooplankton δ15N in the field.

Description: Eggs and larvae of Baltic sprat Sprattus sprattus L. were collected during 14 cruises covering the spawning season in 2002 in Bornholm Basin. Main egg and larval production was in April, with a second small peak in June 2002. The in situ larval abundance was corrected for transport processes by hydrodynamic model runs. Corrected larval abundance estimates were compared to initial larval production to derive an index of larval mortality. This index suggested a much higher survival of summer- over spring-born sprat larvae, with pronounced differences in survival for larvae 〉11 mm. Independent evidence for this survival pattern was gained by measured RNA:DNA ratios in sprat larvae hatched from April to July 2002 and was linked to temporal variability in potential prey abundance. We found higher mean but less variable RNA:DNA ratios in spring- than in summer-born larvae, indicating a strong selection for fast growth in April and May but a less selective environment in June and July. Zooplankton data revealed high naupliar concentrations of Acartia spp. (a key dietary component of sprat) in April and May, but very low concentrations of larger prey items such as copepodites or adults. In contrast, abundance of larger prey increased considerably in June and July. The results suggest that larger sprat (〉11 mm) in April and May 2002 may have been food limited and, therefore, had lower rates of survival, supporting the underlying hypothesis of size-specific, temporally limited ‘windows of survival’ linked to the availability of suitable prey.

Description: The intermediate disturbance hypothesis (IDH) predicts maximal diversity at intermediate levels of disturbance, but the validity of this hypothesis is controversially discussed. In this study, results of a field experiment, which was conducted on the northern-central Chilean coast, are presented. Fouling communities on artificial settlement substrata were studied. A total of 7 disturbance frequencies were applied to previously established communities, and a single disturbance event resulted in a removal of ~20% of the biomass. Species cover was estimated at the end of the experimental period, and it was found that diversity was strongly affected by disturbance frequency. With high disturbance frequencies the composition of the community was changed, with a decrease in the solitary ascidian Pyura chilensis (Molina 1782). The decrease of P. chilensis resulted in an increase of the colonial ascidian Diplosoma sp. A unimodal relationship between disturbance frequency and species richness was found, supporting the IDH. The results suggest that disturbance sustains diversity by reducing the abundance of the dominant species (e.g. P. chilensis), preventing competitive exclusion of the subordinate species, thus allowing subordinate species to re-emerge when competition is alleviated by disturbance. The results also suggest that these species show a trade-off between competitive and colonizing abilities, pointing to the existence of a competitive hierarchy. Therefore, the presence of competitive exclusion and disturbance-induced suppression of the dominant species remains a crucial mechanism, permitting species coexistence in the context of the IDH in the system studied.

Description: We describe an integrated database on European macrobenthic fauna, developed within the framework of the European Network of Excellence MarBEF, and the data and data integration exercise that provided its content. A total of 44 datasets including 465354 distribution records from soft-bottom macrobenthic species were uploaded into the relational MacroBen database, corresponding to 22897 sampled stations from all European seas, and 7203 valid taxa. All taxonomic names were linked to the European Register of Marine Species, which was used as the taxonomic reference to standardise spelling and harmonise synonymy. An interface was created, allowing the user to explore, subselect, export and analyse the data by calculating different indices. Although the sampling techniques and intended use of the datasets varied tremendously, the integrated database proved to be robust, and an important tool for studying and understanding large-scale long-term distributions and abundances of marine benthic life. Crucial in the process was the willingness and the positive data-sharing attitude of the different data contributors. Development of a data policy that is highly aware of sensitivities and ownership issues of data providers was essential in the creation of this goodwill.

Description: Nanoplankton and picoplankton abundance and community grazing on picoplankton were determined in summer and autumn at several stations in a productive coastal environment (Georges Bank, NW Atlantic Ocean) and in an oligotrophic oceanic ecosystem (Sargasso Sea). Ranges of heterotrophic nanoplankton (HNAN) abundance were 1.2 to 3.6 x 103 cells ml-1 on Georges Bank, and 2.2 to 6.8 x 102 cells ml-1 in the Sargasso Sea. Ranges of phototrophic nanoplankton (PNAN) abundance in these ecosystems were 1.9 to 6.0 x 103 and 1.3 to 4.7 x 102, respectively. Mixotrophic nanoplankton (MNAN), operationally defined here as chloroplast-bearing nanoplankton that ingested fluorescent tracers, comprised an average of 12 to 17% of PNAN in surface waters in both environments during August and October. Mixotrophs at specific stations constituted as much as 38% of total PNAN abundance on Georges Bank and 30% in the Sargasso Sea. Mixotrophs represented up to 39% of the total phagotrophic nanoplankton abundance (MNAN/[MNAN + HNAN]). Community grazing impact was estimated from the disappearance of fluorescent prey surrogates (fluorescently labeled bacteria, FLB; cyanobacteria, FLC; and 〈\3 µm algae, FLA). Absolute grazing rates (total picoplankton cells removed d-1) on Georges Bank exceeded those in the Sargasso Sea due to the greater abundances of predators and prey. However, there was overlap in the specific grazing losses at the 2 sites (ranges = 0.08 to 0.38 d-1 in the coastal ocean and 0.05 to 0.24 d-1 in the oligotrophic ocean). Rates of bacterivory were in approximate balance with rates of bacterial production (3H-thymidine uptake), but production exceeded bacterivory on Georges Bank during the summer cruise. These data are among the first documenting the impact of grazing on picoplankton in these environments, and they are consistent with the prediction that nanoplanktonic protists are major predators of picoplankton. While the proportion of phototrophs that are phagotrophic was highly variable, our study indicates that algal mixotrophy is widespread in the marine environment, occurring in both coastal and oligotrophic sites, and should be considered quantitatively in microbial food web investigations.

Description: A cyclonic cold-core eddy in the Northeast Atlantic of about 100 km in diameter at the sea surface was investigated in May 1985, approximately 3 wk after it had separated from the Polar Front. A strong thermocline, which was shallower but more pronounced than in the ambient water, separated a warm surface layer within the eddy from deeper cold water, while horizontal salinity gradients marked the boundary to the ambient water. The cold-core eddy could be distinguished from amblent Northeast Atlantic water in terms of its nutrient chemistry, phytoplankton species distribution and abundance, bacterial numbers and cell size. The surface layer of the eddy was distinct from deeper eddy water, and was characterized by high concentrations of chlorophyll a, total phytoplankton biomass, dinoflagellates and bacteria. At the eddy's margin diatoms were predominant. It is argued that the physical isolation of the eddy surface layer due to the formation of a shallow thermocline led to rapid utilisation of nutrients. This probably enabled the development of a dinoflagellate-dominated phytoplankton population and of organisms capable of heterotrophic regenerative processes.

Description: Bimodal depth distribution patterns observed for sprat Sprattus sprattus larvae in previous field studies conducted in the deep basins of the Baltic Sea have led researchers to hypothesise that larval sprat condition was depth-dependent. We examined this hypothesis by measuring morphological, biochemical and otolith-based proxies for nutritional condition in sprat larvae collected in discrete 5 m depth intervals from the surface to the bottom in the central Bornholm Basin. Similar to earlier studies, larval sprat were most abundant in 2 depth strata (0 to 10 and 65 to 75 m). Their nutritional condition in surface and deep waters was not uniformly expressed by the different indices. For example, sprat larvae from 0 to 10 m could not be distinguished from conspecifics caught at 65 to 75 m by a long-latency condition proxy (otolith-based growth rates). Similarly, a medium-latency proxy (RNA:DNA) did not suggest differences in condition between the depths. However, short-latency proxies (protein:standard length and DNA:dry weight) supported the depth-dependent condition hypothesis. The lack of correspondence and pitfalls associated with the use and interpretation of multiple condition indices (e.g. the influences of temperature and body size) are discussed and recommendations to strengthen these various metrics are provided.

Description: The occurrence of the ctenophore Mnemiopsis leidyi as a new invasive species in the Baltic Sea and the potential consequences for fish stock recruitment was investigated in spring 2007. The study focused on the Bornholm Basin, which serves as the major spawning ground for cod and sprat, the commercially most important fish stocks in the Baltic. The distribution pattern of M. leidyi revealed a substantial overlap with cod eggs. The observed predation of M. leidyi on eggs has the potential to alter the recruitment success of cod, which is the top predator in the system and, thus, to change the Baltic food-web structure.

Description: Deep-sea whale falls create sulfidic habits Supporting chemoautotrophic communities, but microbial processes underlying the formation Of Such habitats remain poorly evaluated. Microbial degradation processes (sulfate reduction, methanogenesis) and biogeochemical gradients were studied in a whale-fall habitat created by a 30 t whale carcass deployed at 1675 m depth for 6 to 7 yr on the California margin. A variety of measurements were conducted including photomosaicking, microsensor measurements, radio-tracer incubations and geochemical analyses. Sediments were Studied at different distances (0 to 9 in) from the whale fall. Highest microbial activities and steepest vertical geochemical gradients were found within 0.5 m of the whale fall, revealing ex situ sulfate reduction and in vitro methanogenesis rates of up to 717 and 99 mmol m(-2) d(-1), respectively. In sediments containing whale biomass, methanogenesis was equivalent to 20 to 30%, of sulfate reduction. During in vitro sediment studies, sulfide and methane were produced within days to weeks after addition of whale biomass, indicating that chemosynthesis is promoted at early stages of the whale fall. Total sulfide production from sediments within 0.5 m of the whale fall was 2.1 +/- 3 and 1.5 +/- 2.1 mol d(-1) in Years 6 and 7, respectively, of which similar to 200 mmol d(-1) were available as free sulfide. Sulfate reduction in bones was much lower, accounting for a total availability of similar to 10 mmol sulfide d(-1). Over periods of at least 7 yr, whale falls can create sulfidic conditions similar to other chemosynthetic habitats Such as cold seeps and hydrothermal vents.

Description: Microchemical otolith analyses have been shown to provide valuable information on the life history, dispersal and stock characteristics of teleost fish. In the present study, the suitability of this technique for identifying the origin and distribution of Atlantic cod Gadus morhua L. from the Baltic Sea was examined using laser ablation-ICPMS. The capacity to distinguish individuals from different Baltic Sea stocks and from the adjacent North Sea stock based on incoporation of stock-specific elemental fingerprints along otolith growth axes was investigated. It was further tested if different origins led to spawning-site specific element concentrations in otolith cores. The results indicate that microchemical analyses of Baltic cod otoliths are applicable for differentiating individuals of different stocks. Analyses of similarities including 12 element/calcium ratios resulted in significant differences between individuals from the eastern and the western Baltic Sea and between North Sea and Baltic Sea samples. Sr/Ca, Ba/Ca, Y/Ca, Mg/Ca, Zr/Ca and Mn/Ca ratios had the strongest discriminatory power. A further separation of individuals caught in 3 different spawning grounds of the eastern Baltic, however, was not possible. Elemental compositions from the core regions of otoliths from young of the year cod caught in eastern and western Baltic Sea spawning grounds showed significant differences in Sr/Ca, Ba/Ca and Mg/Ca concentrations. Analyses of similarities again showed significant differences between these areas for juveniles. This study demonstrates the potential of otolith microchemical analyses to provide important information about the stock structure and connectivity of G. morhua in the Baltic Sea.

Description: Climate change has the potential to profoundly influence the community structure and function of marine ecosystems. Prior to testing the consequences of altered environmental conditions on ecosystem functioning, it is first necessary to better understand how the functioning of an ecosystem is affected by its structure. Using phytoplankton communities with 4 naturally co‑occurring coccolithophores including species of Emiliania, Gephyrocapsa, and Calcidiscus collected off the Azores, we experimentally tested whether varying initial dominance leads to different competitive outcomes and consequently affects community functioning, such as biomass and carbon accumulation. We manipulated initial community structure by creating 5 different dominance scenarios: (1) all species contributing evenly to total initial biomass, and (2–5) one of each species contributing 4× that of the remaining 3 species to total initial biomass. All 4 species were simultaneously grown in monocultures starting with the same total initial biomass as the communities. Monocultures differed significantly in total final biomass, particulate inorganic carbon, and particulate organic carbon content. Priority effects in the communities caused the initially dominant species to remain dominant during the stationary phase in 3 out of 4 cases. However, despite varying dominant species and different outcomes in the monocultures, community functioning was unaffected. We suggest that selective and facilitative effects are responsible for the equalization of community functioning. We conclude that monoculture experiments are not sufficient to predict whole-community responses, since species interactions can significantly alter the expected functional outcome.

Description: Droop’s cell-quota model is the most successful description of phytoplankton growth in laboratory cultures and is increasingly being introduced into the ecosystem components of biogeochemical models. Although the Droop model’s parameters can be easily interpreted in biological terms, it was nevertheless derived empirically and lacks a sound mechanistic foundation. Here we derive Droop’s model from a simple optimality condition which maximises net growth rate. Our approach links the maximum cell quota to the cost of nutrient acquisition and suggests that respiration is influenced more strongly by C fixation than by N assimilation.

Description: The growth and development of the aragonitic CaCO3 otoliths of teleost fish could be vulnerable to processes resulting from ocean acidification. The potential effects of an increase in atmospheric CO2 on the calcification of the otoliths were investigated by rearing Atlantic cod Gadus morhua L. larvae in 3 pCO2 concentrations—control (370 µatm), medium (1800 µatm) and high (4200 µatm)—from March to May 2010. Increased otolith growth was observed in 7 to 46 d post hatch (dph) cod larvae at elevated pCO2 concentrations. The sagittae and lapilli were usually largest in the high pCO2 treatment followed by the medium and control treatments. The greatest difference in mean otolith surface area (normalized to fish length) was for sagittae at 11 dph, with medium and high treatments being 46 and 43% larger than the control group, respectively. There was no significant pCO2 effect on the shape of the otoliths nor were there any trends in the fluctuating asymmetry, defined as the difference between the right and left sides, in relation to the increase in otolith growth from elevated pCO2.

Description: This experimental study investigated the vertical migratory response of benthic forammifera in sediments that were initially oxygenated, to variations in dissolved-oxygen concentrations ranging from well-oxygenated to dysaerobic conditions. Two box cores, with a carpet of polychaete tubes extending above the sediment-seawater interface, were recovered from 71 m water depth in Oslofjord, southern Norway. The seawater oxygen concentration of one box core was decreased every 4 wk, to a minimum value of 〈0.2 ml 0; 1'. Aerated seawater circulation was maintained in the other (control) box core. In a time course, 2 subcores were taken from each mesocosm every 4 wk and sectioned in 0 5 cm mtervals to depths of 2.0 cm, and 1.0 cm intervals from 2.0 to 4.0 cm. The portion of the polychaete tubes extending into the overlymg water was collected and treated as an additional sample. Live foramimferal distributions were examined in the 〉 63 pm fraction of each interval shallower than 2 cm using both an ATP assay and a rose Bengal staining method, while only rose Bengal was used to assess distributions from 2 to 4 cm. Results indicate that larger numbers of live and stained forammifera are found on the polychaete tubes and in the superficial 0.5 cm of sediment after exposure to dysaerobic conditions compared to original and control abundances. After re-oxygenation for 4 mo, the foraminifera re-migrated into the sediments, exhibiting distributions similar to those of the control mesocosm. These observations suggest that some species actively migrate to a microenvironment with a particular oxygen concentration, rather than maintaining a stable position with respect to the sediment-water interface.

Description: The diazotrophic cyanobacterium Trichodesmium is a significant contributor to marine nitrogen and carbon cycles and has been incorporated in biogeochemical ocean circulation models. To date, parameterization of light as a controlling factor for nitrogen fixation has been based on field observations, where factors other than light also affect Trichodesmium physiology. Here we present data on light-dependent (15 to 1100 µmol quanta m–2 s–1) diazotrophic growth from controlled laboratory experiments and their implications for modeling approaches. We supply a simple empirical model to describe nitrogen fixation by Trichodesmium in batch cultures. Diazotrophic growth of axenic Trichodesmium IMS-101 was light saturated at 180 µmol quanta m–2 s–1 and did not vary significantly at higher photon irradiances up to 1100 µmol quanta m–2 s–1 (μcarbon based ≈ 0.26 d–1). Chlorophyll a (chl a) normalized N2 fixation rates were significantly affected by light intensity during mid-exponential growth (0.74 to 4.45 mol N fixed mol chl a–1 h–1) over the range of photon irradiances tested. In contrast, nitrogen fixation rates normalized to the cellular carbon content were relatively unaffected by light intensity (0.42 to 0.59, averaging 0.5 mmol N mol particulate organic carbon [POC]–1 h–1). Trichodesmium carbon biomass can be used to estimate the nitrogen input by this diazotroph into the ocean; the maximum input rate is 350 nmol N fixed l–1 h–1.

Description: Phytoplankton (〉100 µm) abundance was studied in the open waters of the Gulf of Aqaba during the summer stratification period of 1996. A succession took place among the major phytoplankton groups, with diatom numbers decreasing throughout the summer. The diazotrophic cyanobacteria Trichodesmium spp. became more prominent as the stratification period progressed; 5 Trichodesmium species were identified: T. thiebautii, T. erythraeum with tuft-shaped colonies and Trichodesmium sp. with puff-shaped colonies were common at ~102 colonies m-3 throughout the stratification period, whereas T. tenue and T. hildebrandtii were more rare. A bloom of T. thiebautii and T. erythraeum with 〉106 tuft colonies m-3 was observed in coastal waters of the Gulf during fall 1997. Tuft-shaped colonies were dominant near the surface, while puff-shaped colonies of Trichodesmium sp. were mainly found in the bottom half of the photic zone. These depth distributions were maintained for more than 2 mo, suggesting that the 2 colony types occupied distinct niches. Puff-shaped colonies were found to have higher chlorophyll a contents than tufts, but their photosynthetic activities were not significantly different. Fatty acid analysis of dominant plankton species yielded new trophic relationships for Trichodesmium spp. The Trichodesmium spp.-specific fatty acid C22:2 ω6 was found in Macrosetella gracilis (the sole copepod to graze on Trichodesmium spp.) and in chaetognaths, suggesting that these carnivorous zooplankton fed on M. gracilis. Furthermore, this fatty acid was observed in the filter-feeding Salpa maxima, which was abundantly present in the Gulf of Aqaba during June 1997.

Description: Anterior/posterior (a/p) compression of the vertebral column, referred to as 'short tails', is a recurring event in farmed Atlantic salmon. Like other skeletal deformities, the problem usually becomes evident in a late life phase, too late for preventive measures, making it difficult to understand the aetiology of the disease. We use structural, radiological, histological, and mineral analyses to study 'short tail' adult salmon and to demonstrate that the study of adult fish can provide important insights into earlier developmental processes. 'Short tails' display a/p compressed vertebrae throughout the spine, except for the first post-cranial vertebrae. The vertebral number is unaltered, but the intervertebral space is reduced and the vertebrae are shorter. Compressed vertebrae are characterized by an unchanged central part, altered vertebral end plates (straight instead of funnel-shaped), an atypical inward bending of the vertebral edges, and structural alterations in the intervertebral tissue. The spongiosa is unaffected. The growth zones of adjacent vertebrae fuse and blend towards the intervertebral space into chondrogenic tissue. This tissue produces different types of cartilage, replacing the notochord. The correspondence in location of intervertebral cartilage and deformed vertebral end plates, and the clearly delimited, unaltered, central vertebral parts suggest that the a/p compression of vertebral bodies is a late developmental disorder that may be related to a metaplastic shift of osteogenic tissue into chondrogenic tissue in the vertebral growth zone. Given the lack of evidence for infections, metabolic disorders and/or genetic disorders, we propose that an altered mechanical load could have caused the transformation of the bone growth zones and the concomitant replacement of the intervertebral (notochord) tissue by cartilaginous tissues in the 'short tails' studied here. This hypothesis is supported by the role that notochord cells are known to play in spine development and in maintaining the structure of the intervertebral disk.

Description: Multiple stable isotope and fatty acid analyses were applied to examine food web dynamics in an eelgrass Zostera marina L. system in the western Baltic Sea. Samples of eelgrass, epiphytic algae, sand microflora, red algae, phytoplankton and main consumer species were collected in June 2002. delta C-13 values of primary producers ranged from -9.6%. for eelgrass to the most depleted value of -34.9%. for the most abundant red alga, Delesseria sanguinea, Epiphyte delta C-13 (-11.3 parts per thousand.) was close to the value for eelgrass, whereas sand microflora and phytoplankton showed intermediate values (-20.0 and -22.6 parts per thousand, respectively). delta C-13 values of consumer species varied from - 12.2 parts per thousand in the gastropod Lacuna vincta to -23.9 parts per thousand in the amphipod Amphitoe rubricata. Epiphytes, sand microflora and phytoplankton had relatively similar fatty acid signatures, indicating a dominance of diatoms. Fatty acid composition of the main consumers included the biomarker fatty acids for diatoms and red algae, whereas those for eelgrass were negligible. The stable isotope data in combination with the results of the fatty acid analysis strongly indicated that the food web in this eelgrass community was based on epiphytes, sand microflora and red algae. Additionally, the continuous distribution of delta N-15 values implied a food web that was characterized by a large proportion of generalist feeders in every group of consumers and by a high degree of omnivory.

Description: Nitrogen fixation rates and related plankton parameters were determined at 8 stations in the Baltic proper and Mecklenburg Bay in 1997 and 1998. Nitrogen fixation was measured with 15N-tracer method in unenriched samples. Measurable nitrogen fixation rates were found from July to October, with highest rates in August when heterocystous cyanobacteria formed blooms. Nitrogen fixation rates measured during a moderate bloom in 1998 were related to biomass of heterocystous and coccoid (non-heterocystous) cyanobacteria and primary production. The size fraction 〈10 µm contributed significantly to total nitrogen fixation both during day and night. It is discussed whether this may be due to small, non-heterocystous cyanobacteria, which were abundant in summer and autumn. They may separate photosynthesis and nitrogen fixation temporally (day and night) and may be especially responsible for the high nitrogen fixation rates observed in the dark. As the fraction of pico- and nanoplankton was not considered in earlier studies, a new budget of nitrogen fixation in the Baltic proper has been estimated. On average, daily nitrogen fixation rates of 2.5 mmol N m-2 d-1 (in July/August 1997/1998) and mean annual nitrogen fixation of 125 mmol N m-2 yr-1 were estimated for the Baltic proper. The high variability is discussed. For summer 1998, a budget of the nitrogen cycle at the main station in the Gotland Sea is give

Description: Seabirds, like all marine endotherms, have to compensate for the extensive cooling effect of water when diving. Alone among them, cormorants (Phalacrocoracidae) have a wettable plumage and are predicted to require disproportionately large amounts of food to balance heat losses. These piscivorous birds are thus thought to have a detrimental impact on fish stocks. However, we show here that even in great cormorants from Greenland, which dive in water at 3 to 7°C, daily food intake is lower than for well-insulated European seabirds. Despite their wettable plumage, cormorants thus appear to manage their energy budgets in a remarkably efficient way. Nevertheless, the specific foraging strategies which enable this performance make cormorants dependent on high prey density areas, a feature that should be taken into account by future management plans.

Description: In Eckernförde Bay (western Baltic Sea) pockmark structures are induced by groundwater seeping out of the sediment. On 3 occasions in winter and spring 1993-94 we investigated the influence of groundwater on the reduction of salinity, on porewater chemistry, and on bacterial activities (methane oxidation and sulphate reduction). In 2 out of 3 sampling campaigns groundwater discharge could be detected. The concentration gradients of Cl- and SO4= are moved towards the sediment surface by the vertical advection of groundwater during seep times. Without groundwater discharge the porewater chemistry resembled the control site. Compared to the control site, the methane oxidation and sulphate reduction rates were elevated at the pockmark site, reaching maximum values of 49 and 269 µmol l-1 d-1 respectively. The groundwater venting from the pockmark had an end member composition of 80 mM Na+, 1.0 mM Ca++ and was depleted in Mg++. Due to mixing of these major cations along the groundwater/seawater interface, no CaCO3 precipitation was found around the pockmark site.

Description: Living (Rose Bengal stained) foraminifera in gas-hydrate-influenced sediments at the Cascadia convergent margin were investigated. Foraminiferal assemblages from the southern Hydrate Ridge and neighboring basins were compared in terms of abundances, vertical distribution, diversity, and species composition. At Hydrate Ridge, the presence of shallow gas hydrates and increased porewater sulfide concentrations was indicated by extensive bacterial mats of Beggiatoa sp. and clam beds of the bivalve mollusk Calyptogena sp., generating different biological zones. Living foraminifera were found in all biological zones, in sediment layers down to 5 cm. They showed highly variable densities within all zones. The average abundance of benthic foraminifera at Hydrate Ridge differs from neighboring basins. Average species diversities are comparable between biological zones, while the average number of species increases from bacterial mats to clam fields and surrounding sediments. Foraminifera can be characterized by 5 principal component communities which explain 97.3% of the variance of the live assemblages at the southern Hydrate Ridge and neighboring basins. At Hydrate Ridge, 2 foraminiferal zones can be distinguished: (1) an Uvigerina peregrina community which characterizes sediments covered with bacterial mats and clam fields; (2) a ?Spiroplectammina biformis community in the surrounding non-seep sediments. Foraminiferal assemblages in the neighboring Western and Eastern Basin differ from the Hydrate Ridge stations.

Description: Somatic growth and nucleic acid content were studied in North Sea houting Coregonus oxyrhinchus larvae fed exclusively on dry diets at 2 temperatures (8.4 and 17.5°C) during a 32 d rearing experiment. The higher temperature enhanced growth significantly. Mean dry weights at the end of the experiment were 3.6 mg (SD = 1.07, range 1.4 to 5.7 mg) and 31.5 mg (SD = 21.9, range 3.6 to 96.0 mg), and mean standard lengths were 17.7 ± mm (SD = 1.6, range 11.8 to 19.6 mm) and 25.5 mm (SD = 4.2, range 17.0 to 35 mm), respectively. Significant responses to temperature were also found in the nucleic acids. However, these differences were not as remarkable and gave indications that differences in protein growth between treatments was based on protein biosynthesis being driven by the activity of the ribosomes, rather than their number. The use of the degree-day approach to normalize the data clearly showed the temperature-dependence of somatic growth. Only small differences in growth and nucleic acid content were observed in the comparable range of the first 280 degree-days in fed fish. In non-fed fish, the starving potential was very similar (approx. 350 degree-days). Shifts in growth pattern from predominantly hyperplasia to predominantly hypertrophy were detected, with ongoing growth at both temperatures.

Description: The combined and interactive effects of climatic and ecological factors are rarely considered in marine communities. We designed a factorial field experiment to analyze (1) the interactive effects of ambient UV radiation and consumers; and (2) the effects of photosynthetically active radiation (PAR 400 to 700 nm), UVA (320 to 400 nm) and UVB (280 to 320 nm) radiation on a marine hard-bottom community in Nova Scotia, NW Atlantic. Species recruitment and succession on ceramic tiles were followed for 5 mo. We found strong negative UV effects on biomass and cover of the early colonizing macroalga Pilayella littoralis, whereas UVB was more harmful than UVA radiation. Consumers, mainly gammarid amphipods, increased P. littoralis biomass when UV was excluded, probably through fertilization. These initially strong and interacting UV and consumer effects on total biomass and cover diminished as species succession progressed. Species diversity was not affected by experimental treatments, but significant shifts in species composition occurred, especially at the recruitment stage. Red algae were most inhibited by UV, whereas sedentary invertebrates and some brown algae tended to increase under UV exposure. Consumers suppressed green and filamentous brown algae, but favored the other groups. Again, these effects diminished during the later stages of succession. We conclude that UV radiation can be a significant structuring force in early successional benthic communities, and that consumers can mediate its effects.

Description: Small cetaceans are susceptible to incidental mortality in the various forms of gillnet fisheries throughout their range. Research conducted since 1994 has shown that acoustic alarms (pingers) emitting high-frequency pulsed sounds effectively reduce the number of harbor porpoise Phocoena phocoena casualties in sink gillnets. However, the mechanisms behind the effects of pingers were still not understood. Until now, advantages and risks associated with their widespread use could not be evaluated. Here we present the results of 2 field experiments: (1) theodolite-tracking of harbor porpoises exposed to a single PICE-pinger in Clayoquot Sound, Vancouver Island, Canada and (2) herring Clupea harengus capture rates in surface gillnets equipped with and without acoustic alarms (Dukane Netmark 1000, Lien, PICE) in the Baltic Sea herring fishery at Rügen Island, Germany. Our results show that harbor porpoises do not seem to react to an experimental net in their foraging area (n = 172 groups, median group size = 2 porpoises). Porpoise distance from the mid-point of the net was distributed around a median of only 150 m (range 4 to 987 m). A net equipped with an acoustic alarm, however, was avoided (n = 44 groups) within audible range (distance distribution median = 530 m, range 130 to 1140 m). The porpoises were thus effectively excluded from the ensonified area. Herring, one of the main prey species of harbor porpoises, were not affected by the acoustic alarms tested (n = 25407 fish captured). The advantages and risks of using acoustic alarms to mitigate by-catch are discussed.

Description: Pelagic and demersal juvenile Baltic cod Gadus morhua L. were collected on the slope and the top of Rønne bank in the Baltic Sea during 2 cruises in November and December 1998. The objective of this study was to evaluate distinct changes in otolith increment width observed in demersal juveniles by comparison with laboratory-reared individuals, and to investigate the factors determining variation in these increments. The different increment-width patterns were identified with a method based on the widths of consecutive increments. Otolith increment widths of juvenile cod were found to be highly variable within and between individuals, in both the experimental and the field samples. The first change in increment pattern observed in the field samples was related to settling. The formation periodicity of increments within the different pattern intervals was confirmed with a growth model based on otolith growth rates of juvenile cod reared in the laboratory under different conditions. In this model, otolith growth rate was expressed as a function of rearing temperature and fish dry weight. Otolith growth of the field samples was calculated using ambient temperatures obtained from a 3D-circulation model. The best fit to observed otolith growth rates was obtained under the assumption that fish on the slope performed daily vertical migrations between the warm surface layer and the cold bottom layer. The data suggested that fish stayed in the surface layer during the first increment-pattern interval, performed vertical migrations during the second interval, and stayed in association with the seafloor in the subsequent interval, corresponding to the time after the breakdown of the thermocline.

Description: Zooplankton synthesise astaxanthin, a carotenoid pigment believed to protect against high-energy irradiance, from precursors in their diet. Different patterns of astaxanthin vertical distribution would be expected from the benefits of photoprotection, the costs of visual predation and the availability of food. Despite a highly resolved sampling approach (4 m depth intervals), no clear pattern of vertical distribution or correlation with chlorophyll a concentrations was found for Calanus helgolandicus astaxanthin concentrations in the German Bight. This may be attributable to photochromatic adaptation in phytoplankton, diurnal rhythms of copepod grazing and, particularly, vertical migration. A 2 × 2 factorial (light × food) experiment showed that total astaxanthin concentrations in C. helgolandicus were entirely determined by the dynamics of esterified astaxanthin. As expected, the concentrations of astaxanthin esters decreased in starved individuals in the dark, while the combined presence of food and light resulted in an increase of esterified astaxanthin. Similar levels of esterified astaxanthin were maintained in both starving individuals exposed to light and feeding individuals in the dark. The latter implies that astaxanthin esterification is per se a process independent of light, although light exposure enhances it. It is proposed that the function of astaxanthin esters is to generally improve the antioxidant protection of storage lipids, also in situations where photoprotection is not required.

Description: Sinking and sedimentation rates of a natural phytoplankton community were simultaneously measured during the course of a diatom winterkpring bloom in a 13m3 experimental mesocosm. Sinking rate was determined directly in settling columns and was calculated from sediment trap catches. The 2 methods yielded significantly different results. Whole-community as well as speciesspecific sinking rates varied over time. These variations were related to changes of the environmental conditions. Over a 26d study period, a total of 7.5g cm-' was collected in the sediment traps. Viable phytoplankton cells were the primary component of the sedimented matter while zooplankton fecal pellets contributed on average less than 10 %. Assuming the Redfield atomic ratio for the collected material, the amount of carbon which sedimented during the winterkpring bloom could be predicted from pre-bloom nutrient concentrations. The daily sedimentation rate varied considerably over time and displayed a characterisbc pattern. This pattern is evidently a function of both suspended phytoplankton biomass and the temporal variation in whole-community sinking rate.

Description: The nekton community was sampled by a Rectangular Midwater Trawl (RMT 25) over the upper 1000 m of the Scotia Sea dunng January 1991. A total of 81 nekton and micronekton species were collected from 2 sites, one in the oceanlc western Scotia Sea (Stn 1) and the other on the northwestern slope of the South Georgia shelf (Stn 2). Species composition, abundance, biomass and day/night vertical distribution were investigated. Crustaceans were the most important group in terms of species numbers (28 species) followed by mesopelagic fish (24), molluscs (15) and coelenterates (11). Species diversity increased with depth and was higher at Stn 2 (76 species) than at Stn 1 (62 specles). Biomass in the upper 1000 m was considerably higher at Stn 1 (94.6 g wet wt m-2 during daytime, 87 g wet wt m-2 during night) than at Stn 2 (10.2 and 23.7 g wet wt m-2, respectively), mostly due to dense concentrations of the tunicate Salpa thompsoni (41.6 g wet wt m-2 during night). The other main contributors to the high biomass at Stn 1 were coelenterates (28.3 g wet wt m-2 during night) and mesopelagic fish (4.9 g wet wt m-2 during night). Euphausiids (Euphausia triacantha and E. superba) accounted for 1.5 g wet wt m-2 at Stn 2 during night, with E. triacantha the more important of the two (1.4 g wet wt m-2). Except for Bathylagus antarcticus all common mesopelagic fishes showed a marked diurnal vertical migration (i.e. Electrona antarctica, Gymnoscopelus brauen, Krefftichthys anderssoni, Protomyctophum bolini). During daylight they stayed in the core of the Circumpolar Deep Water (CDW, 400 to 800 m) and at night they were mainly distnbuted in the Antarctic Surface Water (ASW, 0 to 400 m). Other species with pronounced vertical migration were the hydromedusa Calycopsis borchgrevinki, the squid Brachioteuthis ?picta, and the euphausiid Euphausia triacantha. The scyphomedusae Atolla wyvillei and Periphylla periphylla and the crustaceans Cyphocaris richardi, Gigantocypris mulleri and Pasiphaea scotiae did not appear to migrate and remained concentrated in the CDW. Spatial variability was analysed by multivariate data analyses (clustering techniques) and related to hydrography. Four main groups, characterised by different nekton communities, were derived: (1) a lower mesopelagic nekton community from the deeper layers of the CDW, apparent at both stations, (2) an upper mesopelagic nekton community from the core of the CDW, apparent at both stations, (3) an epipelagic nekton community from the ASW over the South Georgia slope (Stn 2) and finally (4) an epipelagic nekton community from the ASW of the oceanic Scotia Sea (Stn 1). The performance of the midwater trawl is discussed as it has a substantial impact on the catchability of the nekton. The presented data provide new information on the structure and spatial variability of Antarchc nekton communities and emphasise the geographical and vertical discontinuities between communities.

Description: Embryo survival, larval standard length, yolk-sac area, yolk utilization and resistance to starvation were significantly influenced by the female parent and the interaction between both parents of Baltic Sea cod Gadus morhua in a gamete factorial cross of 3 females and 5 males in all possible combinations. The proportion of variance in embryonic survivorship that was due to maternity during incubation was significant during the first 4 d, but was insignificant during the 5 to 9 d interval. During the 5 to 9 d interval, the male–female interaction was significant. Standard lengths at hatch and 5 d post hatch and specific growth rate of unfed yolk-sac larvae were strongly influenced by the female. Equivalent amounts of variation in larval resistance to starvation were explained by maternity and bi-parental interaction. Paternity alone did not explain a significant amount of variation for the traits or processes examined. The bi-parental interactions reported demonstrate a female’s choice of mate could significantly influence the early life survivorship of Baltic cod offspring.

Description: During the Anglo-German Antarctic expedition in February 1982 macroplankton was collected in the waters off the Antarctic Peninsula. Macroplankton compos~tions at 36 stations were compared and degrees of similarity submitted to hierarchical cluster analysis. Results demonstrate strong spatial heterogeneity, which could be attributed to the different water masses in this region. These dissimilarities demarcate 4 provinces, each characterized by a distinct macroplankton community: (l) The 'Oceanic Community' comprises the stations influenced by the Westwind Drift; oceanic forms are typical (e.g. the hyperiid amphipod Vibilia antarctica, and the polychaete Vanadis antarctica). (2) The 'Bellingshausen Water Community' is influenced by Bellingshausen Sea water and oceanic specles are scarce; large numbers of Antarctic krill Euphausja superba occur but shallow water forms are also abundant. (3) The 'Nentic Community' consists of stations in the shelf water of the southern Bransfield Strait; it is charactenzed by postlarvae of several fish species, and meroplanktonic larvae of benthic forms. Large krill concentrations, however, are also encountered. (4) A 'Transitional Community' exists in environments where various water masses mingle (e.g. at shelf slopes). This community lacks typical forms.

Description: The role of transparent exopolymer particles (TEP) and dissolved organic carbon (DOC) for organic carbon partitioning under different CO2 conditions was examined during a mesocosm experiment with the coccolithophorid Emiliania huxleyi. We designed 9 outdoor enclosures (similar to11 m(3)) to simulate CO2 concentrations of estimated 'Year 2100' (similar to710 ppm CO2), 'present' (similar to410 ppm CO2) and 'glacial' (similar to190 ppm CO2) environments, and fertilized these with nitrate and phosphate to favor bloom development. Our results showed fundamentally different TEP and DOC dynamics during the bloom. In all mesocosms, TEP concentration increased after nutrient exhaustion and accumulated steadily until the end of the study. TEP concentration was closely related to the abundance of E. huxleyi and accounted for an increase in POC concentration of 35 2 % after the onset of nutrient limitation. The production of TEP normalized to the cell Abundance of E. huxleyi was highest in the Year 2100 treatment. In contrast, DOC concentration exhibited considerable short-term fluctuations throughout the study. In all mesocosms, DOC was neither related to the abundance of E. huxleyi nor to TEP concentration. A statistically significant effect of the CO2 treatment on DOC concentration was not determined. However, during the course of the bloom, DOC concentration increased in 2 of the 3 Year 2100 mesocosms and in 1 of the present mesocosms, but in none of the glacial mesocosms. It is suggested that the observed differences between TEP and DOC were determined by their different bioavailability and that a rapid response of the microbial food web may have obscured CO2 effects on DOC production by autotrophic cells.

Description: The effect of methane released from decomposing surficial gas hydrates (SGH) on standing stocks and activities of the small-sized benthic biota (SSBB; i.e. bacteria, fungi, protozoa, and meiobenthic organisms) was studied at about 790 m water depth, at the Hydrate Ridge, Cascadia subduction zone. Presence of SGH and elevated sulfide concentrations in the sediment were indicated by extensive bacterial mats of Beggiatoa sp. and clam fields of the bivalve mollusc Calyptogena sp. Vertical and horizontal distribution patterns of the SSBB biomass were derived from DNA and total adenylate (TA) sediment assays. Potential bacterial exoenzymatic hydrolytic activity was measured using fluorescein-di-acetate (FDA) as substrate. Estimates of chemoautotrophic production of particulate organic carbon (POC.) were determined by 14CO2 uptake incubations. Inventories of chl a and pheopigments were determined as parameters of surface water primary produced POC input. Average SSBB biomass in clam field sediments integrated over the upper 10 cm (765.2 gC m-2, SD 190.1) was 3.6 times higher than in the adjacent control sites (213 gC m-2, SD 125). Average SSBB biomass in bacterial mat sediments, which were almost devoid of eukaryotic organisms 〉 31 µm, was 209 gC m-2 (SD 65). Significant correlations between FDA, DNA and plant pigments imply that productivity of the SSBB at SGH sites is only partially uncoupled from the primary production of the surface water. Areal estimates of autotrophic Corg production at control sites, bacterial mat sites and in clam field sites were 5.7, 59.7 and 190.0 mgC m-2 d-1, respectively. Based on different models predicting vertical POC fluxes from surface water primary production and water depth, these autotrophic POC productions account for 5 to 17% (controls), 35 to 68% (bacterial mats), and 63 to 87% (clam fields) of the bulk POC (sum of allochthonous POC input through the water column and sedimentary autochthonous autotrophic POC production) provided at the various sites. At SGH sites inventories of chl a and pheopigments, integrated over the upper 10 cm of the sediment, were half of that found at the control sites. This might be due to enhanced degradation of phytodetritally associated organic matter. The resulting low molecular weight organic carbon compounds might stimulate and fuel sulfate reduction, which is conducted in a microbial consortium with anaerobic methane consuming archaea. This syntrophic consortium might represent a prominent interface between gas hydrate derived carbon and allochthonous Corg flow. We infer that degradation kinetics of SGH is affected by, e.g., seasonally varying input of allochthonous organic matter.

Description: Sinking velocities of more than 300 Nitzschia closterium aggregates were determined during roller table incubation using digital image analysis. To examine the influence of transparent exopolymer particles (TEP) on aggregate settling speed, 3 experiments with different ratios of TEP to cell volume concentration were conducted. The results showed that, for N. closterium aggregates without TEP, sinking velocity (U) was significantly related to the equivalent spherical diameter (ESD) of the aggregates, yielding U (cm s-1) = 1.89 (ESD, cm)0.55. The higher was the specific TEP content of an aggregate, the lower was the sinking velocity and the less pronounced was the size versus velocity relationship. Excess densities (Δρ) of aggregates were derived from velocity measurements and 3-dimensional fractal dimensions (D3) of aggregates were calculated from scaling properties of Δρ. Values for D3 never exceeded 2 and fit well to values of the 2-dimensional fractal dimension (D2) attained from image analysis.

Description: This study tested whether the extreme scarcity of larger nanophytoplankton and microphytoplankton in the Gulf of Aqaba and in the open northern Red Sea is caused by nutrient limitation or by selective removal by grazers. Samples of near surface phytoplankton were incubated on board under a fully factorial combination of release from grazing pressure and release from nutrient stress. Release from grazing pressure by different size classes was obtained by sieving through 100, 20, and 10 µm size mesh screens. Release from nutrient stress was obtained by enrichment of Si alone and a full enrichment by N, P, Si and trace elements. Growth rates of most phytoplankton taxa showed a strong, positive response to the full nutrient enrichment and a weaker, but significant response to grazer exclusion. Several diatom taxa showed a weak positive response to Si enrichment. Thus, bottom-up control of medium-sized algae appears to be more important than top-down control.

Description: Chromophoric dissolved organic matter (CDOM) represents the optically active fraction of the bulk dissolved organic matter (DOM) pool. Recent evidence pointed towards a microbial source of CDOM in the aquatic environment and led to the proposal that phytoplankton is not a direct source of CDOM, but that heterotrophic bacteria, through reprocessing of DOM of algal origin, are an important source of CDOM. In a recent experiment designed at looking at the effects of elevated pCO2 on blooms of the coccolithophorid alga Emiliania huxleyi, we found that despite the 3 different pCO2 levels tested (190, 414 and 714 ppm), no differences were observed in accumulation of CDOM over the 20 d of incubation. Unlike previous mesocosm experiments where relationships between CDOM accumulation and bacterial abundance have been observed, none was observed here. These results provide some new insights into the apparent lack of effect of pCO2 on CDOM accumulation in surface waters, and question the previously proposed mechanisms and rates of CDOM production in natural phytoplankton blooms.

Description: Increased nutrient loading favors macroalgal blooms in eutrophied coastal ecosystems. The main counteracting factor on this bottom-up support is top-down control by consumers. We asked (1) whether herbivore control on 2 bloom-forming macroalgae in the Baltic Sea varies between different algal life stages, (2) whether herbivores selectively feed on Enteromorpha spp. (Chlorophyceae) thereby supporting dominance of Pilayella littoralis (Phaeophyceae), and (3) whether various herbivore species differ in their effects. In comparative field and laboratory experiments, we analyzed herbivore pressure and selectivity on germling density and adult thalli of Enteromorpha spp. and P. littoralis. In the field, herbivores reduced macroalgal recruitment by 80% within 14 d indicating strong herbivore control at early life stages. Recruits of Enteromorpha spp. were significantly preferred over P. littoralis Adult thalli of both algae showed similar growth rates, but grazing rates were significantly higher on Enteromorpha spp. In laboratory experiments, Idotea chelipes (Isopoda), Littorina saxatilis (Gastropoda) and Gammarus locusta (Amphipoda) strongly reduced germling density, whereas effects of L. littorea were weak. I. chelipes and L. saxatilis significantly preferred germlings of Enteromorpha spp. over P. littoralis. I. chelipes had strong effects on adult Enteromorpha spp. but not on P. littoralis. The effects of G. locusta and L. littorea on both adult algae were smaller and not selective, and L. saxatilis did not feed on adults at all. Different herbivore feeding modes are discussed. We conclude that strong and selective herbivory on juvenile and adult Enteromorpha spp. favors dominance of P. littoralis in the Baltic. An effective control of macroalgal blooms in eutrophied areas can be supported by sustaining high herbivore density and species richness considering the variable and complementary effects of different herbivore species on different algal life stages.

Description: Experiments were carried out on Georges Bank, a productive coastal region in the northwestern sector of the North Atlantic Ocean, and in the oligotrophic western Sargasso Sea to examine the effects of nutrient (inorganic nitrogen and phosphorus) and organic carbon (glucose) additions on bacterial and phytoplankton growth. Four experiments were conducted in each environment. Phytoplankton growth was monitored over a 36 h period by following changes in the concentration of chlorophyll in unfiltered seawater and in seawater prefiltered through 5 μm screening to reduce grazing pressure. Bacterial production was estimated initially and after 24 h using the 3H-thymidine (TdR) method in unfiltered seawater and in 1 μm filtrate. Phytoplankton biomass increased significantly in response to nutrient additions in all but 1 experiment, whereas chlorophyll concentrations remained unchanged or decreased in all of the unamended (control) treatments or treatments supplemented with glucose. Responses of the phytoplankton community were similar for the 〈5 μm and unfiltered treatments. Bacterial production increased after 24 h in all of the treatments on Georges Bank, and there was little effect of nutrient or glucose addition in unfiltered seawater relative to unamended controls. However, glucose addition to the 〈1 μm filtrate caused substantial increases in bacterial production relative to controls and N/P-amended treatments in 2 of the experiments from this environment. Glucose had no stimulatory effect (relative to unamended treatments) in 3 of the 4 Sargasso Sea experiments, and only a marginal effect in the fourth. However, the addition of inorganic nitrogen and phosphorus in the latter ecosystem resulted in higher bacterial production (relative to unamended treatments or glucose addition) in 2 of the experiments with unfiltered seawater, and very large increases in 3 of the experiments with 1 μm filtrate. The magnitude of the changes in bacterial production differed greatly between unfiltered and filtered seawater in both ecosystems, indicating an important role for bacterial grazers in controlling bacterial population growth. The results of this study indicate different nutritional restraints on bacterial production in these contrasting environments.

Description: We investigated the nutritional condition of larval fishes caught in daily ichthyoplankton hauls in the ‘Helgoland Roads’ (54°11.18’N and 07°54.00’E) from February to June 2004. We concentrated on larvae of dab Limanda limanda and lesser sandeel Ammodytes marinus in order to compare early life stages of iteroparous and nearly semelparous fish. We analysed length, weight and RNA:DNA ratios as a proxy for condition of the larvae. The relationship between larval nutritional condition and larval size provided an indication of condition-selective mortality due to a loss of larvae in poor condition from the larger size-classes. For the sandeel, larvae in good condition were present in all size-classes, whereas for the dab maximum larval condition increased with increasing size. Variability in both standard length and condition was high in the 2 species during their planktonic stage. Both species showed good nutritional condition in the early to mid portion of the recruitment period and declines in condition in late April. This was more pronounced in larval dab, which showed a higher dependency on feeding conditions than larval sandeel. Together, these findings indicate a more conservative strategy of early life stages of the nearly semelparous sandeel compared with the iteroparous dab.

Description: During a transition period from oxic to anoxic conditions in the bottom water, rates of sulfate reduction and methane production, methane fluxes, as well as concentration profiles of sulfate, sulfide and methane were measured in sediments at a central site of the Gotland Deep (Stn AL 93, 241 m depth), which is regarded as representative for the deepest part of this basin. During this period from 1993 to 1996 oxic conditions in the bottom water prevailed from spring 1994 until summer 1995 with oxygen concentrations decreasing progressively with time. In the sediments methane production occurred primarily in layers below 1 m depth and flux rates of methane to the sediment surface were characterized by a steep concentration gradient from approx. 5 mM at 4 m depth to values close to 30 μM at the surface, determined by diffusion processes and anaerobic oxidation of methane. Both processes were independent of changes at the sediment surface. Differences in the flux rates of methane between the deeper part with a mean value of 259 μmol m-2 d-1 and the upper layers with a mean of 47.7 μmol m-2 d-1 indicate that a considerable proportion of the methane is oxidized within the anoxic horizon of the sediment (71 to 86% in the layer from 40 to 70 cm). Low rates of methane production found within the top 20 cm of the sediment during periods of oxic bottom water increased after depletion of oxygen and resulted in a clear maximum of the methane concentration in the top 2 cm. Sulfate concentrations declined exponentially from values of 11.5 mM in June 1994 and 8.5 mM in October 1995 at the sediment surface to values of 2.5 mM at 20 cm depth and of less than 0.5 mM at 50 to 60 cm depth. High sulfate reduction rates (150 to 250 nmol cm-3 d-1) in the upper part of the sediment (8 to 13 cm) coincided with maxima of sulfide concentrations. During the time period of this investigation an increase of maximum sulfide concentrations in the sediment from 1 to 10 mM was measured together with decreasing oxygen concentrations in the deep water. At the same time sulfate reduction established a small but distinct maximum at the top layer of the sediment (0 to 2 cm). The relative importance of sulfate reduction and methanogenesis in the carbon budget of the Gotland Deep sediments is calculated on the basis of the actual measurements.

Description: Stable carbon isotope fractionation (ε p) of 7 marine phytoplankton species grown in different irradiance cycles was measured under nutrient-replete conditions at a high light intensity in batch cultures. Compared to experiments under continuous light, all species exhibited a significantly higher instantaneous growth rate (μi), defined as the rate of carbon fixation during the photoperiod, when cultivated at 12:12 h, 16:8 h, or 18:6 h light:dark (L/D) cycles. Isotopic fractionation by the diatoms Skeletonema costatum, Asterionella glacialis, Thalassiosira punctigera, and Coscinodiscus wailesii (Group I) was 4 to 6o/oo lower in a 16:8 h L/D cycle than under continuous light, which we attribute to differences in μi. In contrast, ε p in Phaeodactylum tricornutum, Thalassiosira weissflogii, and in the dinoflagellate Scrippsiella trochoidea (Group II) was largely insensitive to daylength-related differences in instantaneous growth rate. Since other studies have reported growth-rate dependent fractionation under N-limited conditions in P. tricornutum, μi-related effects on fractionation apparently depend on the factor controlling growth rate. We suggest that a general relationship between εi and μi/[CO2,aq] may not exist. For 1 species of each group we tested the effect of variable CO2 concentration, [CO2,aq], on isotopic fractionation. A decrease in [CO2,aq] from ca 26 to 3 µmol kg-1 caused a decrease in ε p by less than 3o/oo. This indicates that variation in μi in response to changes in daylength has a similar or even greater effect on isotopic fractionation than [CO2,aq] in some of the species tested. In both groups ε p tended to be higher in smaller species at comparable growth rates. In 24 and 48 h time series the algal cells became progressively enriched in 13C during the day and the first hours of the dark period, followed by 13C depletion in the 2 h before beginning of the following light period. The daily amplitude of the algal isotopic composition (δ13C), however, was 〈=1.5o/oo, which demonstrates that diurnal variation in δ13C is relatively small.

Description: Recent studies in the marine environment have suggested that the limited phenotypic plasticity of cold-adapted species like Atlantic cod (Gadus morhua L.) will cause distributions to shift toward the poles in response to rising sea temperatures. Some cod stocks are predicted to collapse, but this remains speculative because almost no information is available on the thermal tolerance of cod in its natural environment. We used electronic tags to measure the thermal experience of 384 adult Atlantic cod from eight different stocks found in the NE Atlantic. Over 100,000 days of data were collected in total. The data demonstrate that cod is an adaptable and tolerant species capable of surviving and growing in a wide range of temperate marine climates. Total thermal niche ranged from -1.5°C to 19°C; this range was narrower (1°C to 8°C) during the spawning season. Cod in each of the stocks studied had a thermal niche of approximately 12°C, but latitudinal differences in water temperature meant that cod in the warmer, southern regions experienced three times the degree days (~4000 DD year-1) than individuals from northern regions (~1200 DD year-1). Growth rates increased with temperature, reaching a maximum in those cod with a mean thermal history of between 8°C and 10°C. Our direct observations of habitat occupation suggest that adult cod will be able to tolerate warming seas but that cod populations may still be affected because the effects of marine climate change will impact cod at earlier life-history stages or via indirect effects on prey species.

Description: We assessed the seasonal change in trophic interactions at the mesograzer and small predator level in a temperate eelgrass system. This was done through stable isotope (δ13C and δ15N) and fatty acid analyses of eelgrass Zostera marina, attached epiphytes, sand microflora, the red alga Delesseria sanguinea, and the 4 most common mesograzer and predator species. Sampling took place monthly in the western part of the Baltic Sea during a whole year (April 2002 to February 2003). The trophic importance of microalgae was corroborated in all studied species. Red algae were consumed to a lesser extent and eelgrass was of minor importance. The degree of dependence on the main carbon sources was species-specific and varied with time. The small gastropod, Rissoa membranacea, mostly grazed epiphytes (73% on average), whereas the fractions of epiphytes and sand microflora showed a more balanced pattern in the other mesograzer species. Stable carbon data and fatty acid composition strongly suggested that epiphytic algae were the primary source of organic matter for mesograzers in late spring and autumn. In summer, sand microflora were of greater importance as carbon sources, except for the amphipod Gammarus oceanicus for which red algae was the most important. Stable nitrogen values indicated that the degree of carnivory was size-dependent in both omnivorous crustacean species studied; larger individuals generally occupied a higher trophic position than did smaller ones. Furthermore, the isopod Idotea baltica was more herbivorous in summer than in the other seasons. Our results confirm the importance of species-specific and temporal variability for the effect of mesograzers in eelgrass systems. The significance of mesograzers as determinants of food-web structure via the reduction of epiphytes is further corroborated by our study, and the observed high plasticity of mesograzers concerning food sources may promote the stability of eelgrass food webs despite strong seasonal variations in the biomass of primary producers.

Description: Culturing experiments were performed with the benthic foraminifer Ammonia aomoriensis from Flensburg Fjord, western Baltic Sea. The experiments simulated a projected rise in atmospheric CO2 concentrations. We exposed specimens to 5 seawater pCO2 levels ranging from 618 µatm (pH 7.9) to 3130 µatm (pH 7.2) for 6 wk. Growth rates and mortality differed significantly among pCO2 treatments. The highest increase of mean test diameter (19%) was observed at 618 µatm. At partial pressures 〉1829 µatm, the mean test diameter was observed to decrease, by up to 22% at 3130 µatm. At pCO2 levels of 618 and 751 µatm, A. aomoriensis tests were found intact after the experiment. The outer chambers of specimens incubated at 929 and 1829 µatm were severely damaged by corrosion. Visual inspection of specimens incubated at 3130 µatm revealed wall dissolution of all outer chambers, only their inner organic lining stayed intact. Our results demonstrate that pCO2 values of ≥929 µatm in Baltic Sea waters cause reduced growth of A. aomoriensis and lead to shell dissolution. The bottom waters in Flensburg Fjord and adjacent areas regularly experience pCO2 levels in this range during summer and fall. Increasing atmospheric CO2 concentrations are likely to extend and intensify these periods of undersaturation. This may eventually slow down calcification in A. aomoriensis to the extent that net carbonate precipitation terminates. The possible disappearance of this species from the Baltic Sea and other areas prone to seasonal undersaturation would likely cause significant shifts in shallow-water benthic ecosystems in the near future.

Description: The causes and consequences of the coexistence of a number of species in a given habitat have attracted ecological research for several decades. Numerous theories have been developed in this context, including the ‘intermediate disturbance hypothesis’ (IDH), but supportive experimental evidence remains scarce and contradictory, leading scientists to propose the existence of an interaction between disturbance and productivity. This study assessed the interactive effects of disturbance and nutrient enrichment at 1 oligotrophic (Arraial do Cabo) and 1 eutrophic (Guanabara Bay) site on the coast of Brazil. At each site, an epibenthic assemblage was allowed to establish on settlement panels (PVC) for 3 mo prior to a 6 mo manipulation phase comprising nutrient enrichment and disturbance (biomass removal). Our results revealed site-specific diversity-driving processes in the absence of disturbance. Nevertheless, diversity and species richness peaked at both sites at some intermediate level of disturbance, corroborating the IDH. Nutrient enrichment increased total species richness and algal species richness in particular, but only at the oligotrophic site. Only here, did nutrient enrichment eliminate the unimodal species richness pattern observed along the disturbance gradient under ambient nutrient concentrations. Such interactive effects of disturbance and productivity on diversity confirm the general predictions of advanced IDH models, e.g. the Kondoh model. The present study indicates that interactive effects of ‘bottom-up’ and ‘top-down’ processes may explain more of the variation in community diversity than the separate models of disturbance–diversity and productivity–diversity relationships.

Description: Sea turtles are diving ectotherms that are influenced by the temperature of the ambient water, although swimming activity can temper this influence via increased body temperatures enhanced by the thermal inertia of these large animals. We successfully equipped 3 nesting hawksbill turtles Eretmochelys imbricata with time–depth recorders (TDRs) to monitor water temperature and dive depth over the duration of the re-migration interval between 2 successive nesting seasons. Data sets for up to 22 mo were obtained, showing fluctuations in water temperature over the seasons. Nocturnal dive behaviour of the turtles at their foraging grounds revealed an increase in dive duration with decreasing water temperatures in winter. A model is provided to estimate dive duration for the range of temperatures experienced by this species in the wild. The data on vertical velocity during ascent and descent phases as a parameter for activity failed to show thermal dependence. It is concluded that changes in water temperature have an effect on the behavioural ecology of Hawksbill Turtles.

Description: As one of the elements for a model on the food requirements of Humboldt penguins Spheniscus humboldti we determined, via gas respirometry, metabolic rates while swimming and resting in water. During rest in water at 19°C Humboldt penguins (mean body mass 3.6 kg) required 5.95 W kg-1. This corresponds to a thermal conductance in water of 0.2975 W (kg °C)-1 (at Ta 19°C and assuming a Tb of 39°C). When swimming in a 20 m long channel, metabolism rose from 8 W kg-1 at a speed of 0.6 ms-1 to 23.1 W kg-1 at 2.2 m s-1. Transport costs (the cost to move 1 kg of body mass over a distance of 1 m) reached a minimum at 1.4 ms-1 with 8.1 J (kg m)-1, which corresponds to 0.89 J (Nm)-1. We corrected for acceleration and deceleration in the channel to determine transport costs of free-ranging Humboldt penguins travelling at sea, which were calculated as 7 J (kg m)-1 (0.71 J [Nm]-1), at 1.7 m s-1. Birds feeding chicks need to balance the costs of either (1) returning to the breeding island for the night and travelling back to the feeding grounds in the morning or (2) incurring increased thermoregulatory costs associated with resting at sea overnight. Simple calculations show that at water temperatures of 19°C we expect Humboldt penguins to show a tendency to remain at sea overnight if foraging areas are 〉4 km from their island. In colder waters (12°C), this distance increases to 〉9 km. Using previously published data on at-sea activity of Humboldt penguins, we found that foraging costs during chick rearing amount to 340 g anchovies d-1. Finally, we present a general model to convert Humboldt penguin activity data at sea to food requirements.

Description: Latitudinal clines in species diversity in limnic and terrestrial habitats have been noted for well over a century and are consistent across many taxonomic groups. However, studies in marine systems over the past 2 to 3 decades have yielded equivocal results. We conducted initial analyses of the MarBEF (EU Network of Excellence for Marine Biodiversity and Ecosystem Function) database to test for trends in local and regional diversity over the latitudinal extent of European continental-shelf waters (36° to 81°N). Soft-sediment benthic macrofauna exhibit little evidence of a latitudinal cline in local (α-) diversity measures. Relationships with water depth were relatively strong and complex. Statistically significant latitudinal trends were small and positive, suggesting a modest increase in diversity with latitude once water-depth covariates were removed. These results are consistent regardless of whether subsets of the database were used, replicates were pooled, or component taxonomical groups were evaluated separately. Local and regional diversity measures were significantly and positively correlated. Scientific cooperation through data-sharing is a powerful tool with which to address fundamental ecological and evolutionary questions relating to large-scale patterns and processes.

Description: The large temporal and spatial variability in carbon isotope fractionation of marine phytoplankton (ε p) is thought to reflect differences in environmental conditions. Meaningful interpretation of this variability requires an understanding of the processes responsible for phytoplankton isotope fractionation. While numerous factors have been suggested to potentially influence ε p, recent theoretical and experimental evidence has emphasized the primary role of phytoplankton growth rate (µ) and CO2 concentration ([CO2aq]) in controlling ε p. Experimental examination of the relationship of ε p with µ and [CO2aq] in studies using different experimental approaches, however, has yielded inconsistent results. Here we directly compare new and previously published data on ε p as a function of CO2 concentration and growth rate for the marine diatom Phaeodactylum tricornutum. When grown under nitrogen-deficient conditions (nitrate-limited chemostat), ε p of P. tricornutum decreases with increasing growth rate. In contrast, under N-replete conditions ε p values are considerably lower at comparable growth rates and CO2 concentrations and are largely insensitive to a 3-fold increase in growth rate due to increasing photon flux density. In both experimental approaches, ε p shows a relatively small CO2 sensitivity in the range of CO2 concentrations naturally occurring in the ocean (8 to 25 µmol kg-1). Below ca 5 µmol CO2 kg-1, a strong decline in ε p with decreasing [CO2aq] is observed. The apparent difference in ε p responses between nitrate-limited and light-controlled cultures of P. tricornutum suggests a principal difference in carbon acquisition for different growth-rate-limiting resources. A mechanistic explanation is proposed and potential implications for the interpretation of phytoplankton carbon isotope fractionation are discussed.

Description: General concepts of reproductive traits in pelagic marine copepods have been developed largely without data from the microcopepod family Oncaeidae. For a very abundant oncaeid species in the Red Sea, Oncaea bispinosa Böttger-Schnack, reproductive parameters are provided and used for testing how the copepod family Oncaeidae may fit into these concepts. Total number and vertical distribution of eggs of this species have been calculated for populations in the northernmost Red Sea and the Gulf of Aqaba during spring, based on counts of detached egg sacs retained in fine mesh net samples. Reproduction is indicated to take place in the meso- and bathypelagic zones, with maximum abundances of eggs occurring within the deep population centres of females at 250 to 450 m depth. The epipelagic population appeared to be non-reproductive. By calculating the biomass of the eggs and the females using published data on size/weight relationships, a first estimate of the reproductive potential of O. bispinosa was obtained and compared with various other oncaeid species. The presently available data for Oncaeidae in total suggest a relationship between single egg (Ec) and female (Fc) carbon weight (Ec = 0.012 × Fc0.48) that differs from corresponding regressions reported in the literature for other marine pelagic copepods. In addition, regressions are given for minimum and maximum clutch carbon related to female carbon weight. Potential egg production rates of O. bispinosa have been calculated from the observed fractions of egg-bearing females and published data on egg developmental times, as available for larger oncaeids. These production rates appear to be lower than have been observed in experimental studies for different pelagic microcopepod taxa. Remaining uncertainties in generalizing concepts for the scaling of fecundity in pelagic marine copepods are discussed, and the need for more comparable data, especially for the numerically abundant oncaeids, is emphasized.

Description: The sediment beneath and at various distances from the fish farm cages in Cephalonia bay (Eastern Mediterranean) was investigated seasonally through sediment profiling imagery (SPI) as well as through monitoring of geochemical variables and macrofaunal assemblages. The SPI images (SPI) repeatedly showed the same qualitative pattern along the benthic enrichment gradient with readily identifiable attributes such as depth of dark sediment, signs of outgassing and bioturbation marks. Quantitative comparisons showed that a large number of SPI attributes showed significant positive or negative correlation with geochemical and biological attributes describing the effect of fish farming on the seabed. All multivariate patterns obtained through the analysis of SPI attributes were highly correlated to those obtained from standard multivariate analysis of macrofauna during the respective seasons. It is argued that SPI provides an integrated assessment of the sedimentary conditions and therefore may be used as a complement of or even a substitute for standard sampling methods when mapping the effects of aquaculture on silty substrates

Description: This study examines whether or not biogeographical and/or managerial divisions across the European seas can be validated using soft-bottom macrobenthic community data. The faunal groups used were: all macrobenthos groups, polychaetes, molluscs, crustaceans, echinoderms, sipunculans and the last 5 groups combined. In order to test the discriminating power of these groups, 3 criteria were used: (1) proximity, which refers to the expected closer faunal resemblance of adjacent areas relative to more distant ones; (2) randomness, which in the present context is a measure of the degree to which the inventories of the various sectors, provinces or regions may in each case be considered as a random sample of the inventory of the next largest province or region in a hierarchy of geographic scales; and (3) differentiation, which provides a measure of the uniqueness of the pattern. Results show that only polychaetes fulfill all 3 criteria and that the only marine biogeographic system supported by the analyses is the one proposed by Longhurst (1998). Energy fluxes and other interactions between the planktonic and benthic domains, acting over evolutionary time scales, can be associated with the multivariate pattern derived from the macrobenthos datasets. Third-stage multidimensional scaling ordination reveals that polychaetes produce a unique pattern when all systems are under consideration. Average island distance from the nearest coast, number of islands and the island surface area were the geographic variables best correlated with the community patterns produced by polychaetes. Biogeographic patterns suggest a vicariance model dominating over the founder-dispersal model except for the semi-closed regional seas, where a model substantially modified from the second option could be supported.

Description: For a 6 wk period covering the time before, during, and after the phytoplankton spring bloom, macroscopic aggregates (2 0.5 mm diameter) were repeatedly collected and water column properties simultaneously measured at a fixed station in the Southern North Sea. Distinct changes in aggregate structure and composition were observed during the study. Predominantly detrital aggregates during the early phase of the study were followed by diatom-dominated algal flocs around the peak of the bloom. Mucus-rich aggregates containing both algal and detrital components and with large numbers of attached bacteria dominated the post-bloom interval. The phytoplankton succession within the aggregates closely reflected the succession in the water column with a time delay of a few days. Algal flocculation did not occur as a simultaneous aggregation of the entire phytoplankton community, but as a successional aggregat~on of selected diatom species. Although the concentrations of inorganic nutrients diminished considerably during the development of the phytoplankton bloom, the termination of the bloom appeared to be mostly controlled by physical coagulation processes. The importance of biologically-controlled factors for physical coagulation is discussed.

Description: Weight-specific growth rate (G) and growth performance (the fraction of maximum growth realized, G pf) are key demographic characteristics. The ratio of RNA/DNA (RD) can provide information on both G and G pf. Estimating G from RD in larval fish requires an adjustment for the activity of RNA at different temperatures. Based on a meta-analysis of published data, we present a general model for the relationship between G in marine fish larvae and fluorometrically derived RD and temperature (T), and suggest that this model can be used to estimate G in marine fish larvae. Several options for estimating G pf are also considered, including the use of a reference growth rate (Gref). RDs of well-fed larvae appeared to be independent of water temperatures between 4 and 28°C, suggesting that any increase in growth rate with temperature was accomplished by increased activity rather than increased concentrations of RNA. However, for the best-fit meta-analysis RD–T–G model, the relationship between RD and G pf was temperature dependent for fish less than fully fed.

Description: The present study quantifies the spatial heterogeneity of the environmental conditions associated with successful spawning by cod Gadus morhua in the Bornholm Basin. Quarterly means from 1989 to 2003 of the thickness of reproductive volume that enabled egg survival indicate that most favourable spawning conditions were located in the central area, inside the 80 m isobath. On average, spatial patterns were similar in the second and third quarters, but with overall lower thicknesses of reproductive volume and less horizontal extension of conditions suitable for egg survival in the third quarter. The observed basin-wide variation in thickness of reproductive volume and oxygen content inside this volume can result in marked horizontal differences in oxygen-related egg mortality, especially during stagnation years. The spawning habitat selected by adult Baltic cod in the Bornholm Basin was characterised by comparing data on egg abundance with environmental variables measured concurrently with egg collection. A clear preference for spawning at locations in the deep basin is evident after both inflow events, as well as for stagnation periods. In consequence, the closed area for fishing implemented in the Bornholm Basin during the main spawning periods from 1995 to 2003, although located in the northern centre of the basin, did not necessarily ensure undisturbed spawning in stagnation years.

Description: At Hydrate Ridge (HR), Cascadia convergent margin, surface sediments contain massive gas hydrates formed from methane that ascends together with fluids along faults from deeper reservoirs. Anaerobic oxidation of methane (AOM), mediated by a microbial consortium of archaea and sulfate-reducing bacteria, generates high concentrations of hydrogen sulfide in the surface sediments. The production of sulfide supports chemosynthetic communities that gain energy from sulfide oxidation. Depending on fluid flow, the surface communities are dominated either by the filamentous sulfur bacteria Beggiatoa (high advective flow), the clam Calyptogena (low advective flow), or the bivalve Acharax (diffusive flow). We analyzed surface sediments (0 to 10 cm) populated by chemosynthetic communities for AOM, sulfate reduction (SR) and the distribution of the microbial consortium mediating AOM. Highest AOM rates were found at the Beggiatoa field with an average rate of 99 mmol m-2 d-1 integrated over 0 to 10 cm. These rates are among the highest AOM rates ever observed in methane-bearing marine sediments. At the Calyptogena field, AOM rates were lower (56 mmol m-2 d-1). At the Acharax field, methane oxidation was extremely low (2.1 mmol m-2 d-1) and was probably due to aerobic methane oxidation. SR was fueled largely by methane at flow-impacted sites, but exceeded AOM in some cases, most likely due to sediment heterogeneity. At the Acharax field, SR was decoupled from methane oxidation and showed low activity. Aggregates of the AOM consortium were abundant at the fluid-impacted sites (between 5.1 × 1012 and 7.9 × 1012 aggregates m-2) but showed low numbers at the Acharax field (0.4 × 1012 aggregates m-2). A transport-reaction model was applied to estimate AOM at Beggiatoa fields. The model agreed with the measured depth integrated AOM rates and the vertical distribution. AOM represents an important methane sink in the surface sediments of HR, consuming between 50 and 100% of the methane transported by advection.

Description: The present study investigated the effect of temperature on male and female swimming activity of the calanoid copepod Temora longicornis, sampled during winter (February) and summer (August) in the English Channel coastal ecosystem. Video recordings were conducted at 3 temperatures representative of those to which these organisms are normally exposed (13, 16 and 20 degrees C) and one extreme-event temperature (24 degrees C). Examinations of instantaneous velocity and symbolic analysis (i.e. dynamics of swimming states discretized from time series of instantaneous velocity) showed that T. longicornis changed its behaviour when confronted with environmental temperature variations. Swimming speed increased as temperature increased. In warmer water, this copepod displayed higher swimming activity, break periods were less frequent, and the frequency of jumps increased. This phenomenon was amplified when the environmental temperature was increased to 24 degrees C. These observations revealed a considerable tolerance to high temperatures and an ability to adjust to environmental temperature changes. The 'summer population' was less active in the low temperature range, but the swimming speed reached a higher value at higher temperatures than that shown by the 'winter population'. The results of the present study highlighted changes in the individual behaviour of this copepod in response to changing seasonal conditions in the form of swimming activity, and thus its ability to maintain biological processes throughout the year, even in a restrictive environment.

Description: Sediment incubation microcosms, multitrap apparatus and water column variables have been employed to describe the dynamic changes in benthic-pelagic coupling between nutrient pools in the Pontevedra ria, NW Spain, during spring and summer 1998. A comparison of the chemical characteristics of suspended and bed sediment together with sediment nutrient effluxes revealed that particulate organic nitrogen and carbon were progressively depleted upon transit through the nutrient pools. The main fate of particulate organic nitrogen reaching the bed sediment is denitrification, although resuspension is also important. An estimate of net denitrification (Dnet) was made at 2 muddy sites in the ria with a mass balance at the benthic boundary layer. First-order approximations calculate Dnet to be 178 and 182 µmolN m-2 h-1 at the 2 stations, and agree well with previous modelling estimates. Denitrification is highest when upwelling relaxes and the flux of organic matter to the sediment increases. Regular inputs of offshore water ensure water renewal and re-oxygenation of bottom waters, thus preventing anoxia, particularly in the summer. With upwelling, large quantities of ammonium are effluxed to the water column (250 µmolNH4+ m-2 h-1), probably as a result of the bed sediment resuspension engendered by upwelling and stirring of phytodetrital fluff held in suspension as neutrally buoyant material above the sediment surface. We hypothesise that hydrodynamical processes play an important role in determining the quantity of nutrients remineralised in the Pontevedra ria and, in the case of nitrogen, the rate of denitrification at the benthic boundary layer.

Description: Differences with respect to anti-herbivore defense were investigated in invasive and native populations of the seaweed Gracilaria vermiculophylla. Specimens from 6 native populations in East Asia and from 8 populations invasive in Europe and the Mexican Pacific coast were maintained under identical conditions and offered to herbivorous snails from both the native range (Littorina brevicula) and Europe (L. littorea) in no-choice feeding assays. L. brevicula consumed in total significantly larger amounts of G. vermiculophylla tissue than did L. littorea. Further, both snail species least consumed the seaweed specimens originating from either non-native populations or from populations native to the Korean East Sea/Sea of Japan. The Korean East Sea/Sea of Japan had previously been identified as putative donor region of all the invasive populations of G. vermiculophylla. Thus, populations in the donor region as well as non-native populations in different invaded realms feature an increased capacity to resist feeding pressure. Differences in nutrient content did not account for the observed patterns of consumption, as palatability and carbon to nitrogen (C:N) ratio were not significantly correlated. Thus, mechanical or chemical defenses or the content of feeding cues influenced the behavior of the snails. We suggest that low palatability contributed to the invasion success of the species.

Description: The conspicuous retreat of the key species Fucus vesiculosus from the deeper parts of its former distribution area in the Baltic Sea has triggered extensive research on the factors that control its growth. Based on recently obtained knowledge on a large number of potential drivers, we developed a numerical model incorporating effects of abiotic factors on the physiological functions of photosynthesis, respiration, and reproduction and the ecological processes of competition, grazing, and epibiosis. For all input combinations, the model delivers the monthly net growth rate near the bladder wrack’s depth limit and the maximum depth of its vertical distribution. The use of data corresponding to conditions presently observed in the western Baltic Sea sets the year’s maximum algal net growth rate in late spring and 2 minima in early spring and autumn. The depth limit of the wrack’s distribution is set at ~9 m. Light and its absorption by phytoplankton represent by far the most important factors controlling the modeled net growth rate and depth penetration, with the role of epibiosis requiring further investigation. Lacking findings on population dynamics and biotic interactions restrict the generated model to an exploratory rather than a predictive tool.

Description: The impact of seawater acidification on calcifying organisms varies at the species level. If the impact differs between predator and prey in strength and/or sign, trophic interactions may be altered. In the present study, we investigated the impact of 3 different seawater pCO2 levels (650, 1250 and 3500 µatm) on the acid–base status or the growth of 2 predatory species, the common sea star Asterias rubens and the shore crab Carcinus maenas, and tested whether the quantity or size of prey consumed is affected. We exposed both the predators and their prey, the blue mussel Mytilus edulis, over a time span of 10 wk and subsequently performed feeding experiments. Intermediate acidification levels had no significant effect on growth or consumption in either predator species. The highest acidification level reduced feeding and growth rates in sea stars by 56%, while in crabs a 41% decrease in consumption rates of mussels could be demonstrated over the 10 wk experimental period but not in the subsequent shorter feeding assays. Because only a few crabs moulted in the experiment, acidification effects on crab growth could not be investigated. Active extracellular pH compensation by means of bicarbonate accumulation was observed in C. maenas, whereas the coelomic fluid pH in A. rubens remained uncompensated. Acidification did not provoke a measurable shift in prey size preferred by either predator. Mussels exposed to elevated pCO2 were preferred by previously untreated A. rubens but not by C. maenas. The observed effects on species interactions were weak even at the high acidification levels expected in the future in marginal marine habitats such as the Baltic Sea. Our results indicate that when stress effects are similar (and weak) on interacting species, biotic interactions may remain unaffected.

Description: Pseudomonas strains were shown to be regularly associated with the brown macroalga Saccharina latissima from the Baltic Sea, studied over several years, and were identified as producers of the antimicrobially active compound 2,4-diacetylphloroglucinol. These findings support the assumption of a stable association between the Pseudomonas spp. strains and S. latissima in the Baltic Sea. The metabolite profile of the Pseudomonas spp. comprised monoacetylphloroglucinol, 2,4-diacetylphloroglucinol, pyoluteorin and several rhizoxins, which exhibited broad-spectrum antibiotic activities against Gram-positive and Gram-negative bacteria as well as against fungi. Because the antibiotic activities included the inhibition of the 2 algal pathogens Pseudoalteromonas elyakovii and Algicola bacteriolytica, we propose a beneficial effect of these marine pseudomonads on their host S. latissima.

Description: Phytoplankton experience strong and abrupt variations in light intensity. How cells cope with these changes influences their competitiveness in a highly dynamical environment. While a considerable amount of work has focused on photoacclimation, it is still unknown whether processes specific of phytoplankton groups (e.g. calcification and silicification) influence their response to changing light. Here we show that the diatom Phaeodactylum tricornutum and the coccolithophore Emiliania huxleyi respond to an abrupt increase in irradiance by increasing carbon fixation rates, decreasing light absorption through the decrease of light-harvesting pigments and increasing energy dissipation through the xanthophyll cycle. In addition, E. huxleyi rapidly increases calcium carbonate precipitation in response to elevated light intensity, thereby providing an additional sink for excess energy. Differences between the 2 species also emerge with regard to the magnitude and timing of their individual responses. While E. huxleyi show a pronounced decrease in chlorophyll a and fucoxanthin cellular contents following increased light intensity, P. tricornutum has a faster increase in diadinoxanthin quota, a slower decrease in Fv/Fm (ratio of variable to maximum fluorescence) and a stronger increase in organic carbon fixation rate during the first 10 min. Our findings provide further evidence of species-specific responses to abrupt changes in light intensity, which may partly depend on the phytoplankton functional groups, with coccolithophores having a supplementary path (calcification) for the rapid dissipation of excess energy produced after an abrupt increase in light intensity. These differences might influence competition between coexisting species and may therefore have consequences at the community level.

Description: Rising atmospheric CO2 concentrations will have profound effects on atmospheric and hydrographic processes, which will ultimately modify the supply and chemistry of trace metals in the ocean. In addition to an increase in sea surface temperatures, higher CO2 also results in a decrease of seawater pH, known as ocean acidification, with implications for inorganic trace metal chemistry. Furthermore, direct or indirect effects of ocean acidification and ocean warming on marine biota will also affect trace metal biogeochemistry via alteration of biological trace metal uptake rates and metal binding to organic ligands. Currently, we still lack a holistic understanding of the impacts of decreasing seawater pH and rinsing temperatures on different trace metals and marine biota, which complicates projections into the future. Here, we outline how ocean acidification and ocean warming will influence the inputs and cycling of Fe and other biologically relevant trace metals globally, and regionally in high and low latitudes of the future ocean, discuss uncertainties, and highlight essential future research fields.

Description: Three bioassay experiments were performed to study the effects of nutrient and Saharan dust additions on natural diazotrophic communities in the tropical North Atlantic Ocean. Samples for nucleic acid analysis were collected at the beginning and end of 48 h incubations. TaqMan probes specific to 7 diazotrophic phylotypes, viz. filamentous cyanobacteria (Trichodesmium spp.), unicellular cyanobacterial (UCYN) Groups A, B, and C, Gamma A and P Proteobacteria, and Cluster III, were used to quantify nifH DNA abundances. N-2 fixation rates were measured in the same experiments using the N-15(2) gas bubble injection method. N-2 fixation was co-limited by P and Fe. Total nifH abundances increased relative to the control with additions of either Fe or P or both in combination. Additions of dissolved N, alone or in combination with phosphate, induced increases in UCYN-A and Gamma A nifH compared with the control. Saharan dust additions significantly stimulated fixation rates. Abundances of all cyanobacterial and Gamma A nifH phylotypes at least doubled after Saharan dust additions where surface water dissolved Fe concentrations were 〈2 nmol l(-1). Laboratory experiments with cultures of T. erythraeum demonstrated that dust addition promoted colony formation and the persistence of T. erythraeum biomass relative to cultures to which no Fe was added. Our results with both field and laboratory experiments indicate that Saharan dust positively affects diazotrophic phylotype abundances and changes T. erythraeum colony morphology.

Description: Repeated invasions of European waters by the ctenophore Mnemiopsis leidyi offer a unique opportunity to study population dynamics and dispersal in gelatinous zooplankton. Here we followed population establishment in two recently invaded areas, the North and Baltic Sea, and analysed changes in population structure during a 3-yr interval using 7 highly polymorphic microsatellites (representing 191 alleles). A second goal was to reconstruct routes of recent invasive range expansion into the Mediterranean Sea During the study period (2008-2010) populations in North Sea and Western Baltic Sea maintained their allelic composition with virtually unchanged levels of genetic diversity and between-population differentiation. This demonstrates that gene flow between the two regions was limited and indicates successful reproduction in both areas. In contrast, at the eastern distribution limit in the central Baltic (Bornholm Basin) the same measures fluctuated between years and genetic diversity decreased from 2008-2010. In concordance with prior ecological observations, this supports the view that here M. leidyi formed a sink population. In the area of recent range expansion (Mediterranean Sea) we observed high population differentiation for a holoplanktonic species. Among Mediterranean samples collected at sites in Spain, France and Israel pairwise differentiation was between Fst = 0.04-0.16. Despite such differentiation, Bayesian clustering and phylogeographic analysis support the hypothesis that all Mediterranean M. leidyi result from a secondary introduction originating in the Black Sea. Our study contributes to growing evidence that multiple invasions of the same species can vary in their degree of genetic diversity and demonstrates how genetic markers can help to resolve whether gelatinous plankton species form self-sustaining populations.

Description: Climate models predict increases in frequency of summer heat waves. In Europe, such events have already caused declines in seagrass meadows, highlighting the importance of short-term responses of local communities to climate stress. Understanding the variability among populations along the European thermal gradient in response to heat waves is crucial for seagrass conservation and management. Using a mesocosm we compared effects of a simulated heat wave on the photophysiology of Zostera marina populations coming from low (43° N, Adriatic Sea) and high latitudes (56° N, North and Baltic Seas). Measurements before, during and up to 4 wk after the heat wave included photophysiological parameters derived from light response curves generated by PAM fluorometry and gene expression using qRT-PCR. In all 3 populations, initial exposures to thermal stress were characterized by increases in dark adapted effective quantum yield (Y0), maximum electron transfer rate of PSII (ETRmax) and slope of the light response curve (α), coinciding with upregulations of the gene superoxidase dismutase [Mn]. With continuation of the heat wave these initial effects disappeared, demonstrated by declines in Y0, ETRmax and α relative to controls. Z. marina from the Adriatic suffered from the simulated heat wave as much as its high-latitude counterparts. However, we also demonstrate slight photophysiological differences between the populations during the recovery phase, where performance of high-latitude populations continued declining even after water temperatures returned to control levels, while photochemical activity fully recovered in the Adriatic population. These results might draw the attention of future studies and seagrass conservation efforts.

Description: The carbon and nitrogen content of transparent exopolymer particles (TEP) was determined and related to the concentration of TEP as quantified by a colorimetrical method. TEP were produced in the laboratory from dissolved precursors by laminar or turbulent shear. Dissolved precursors were obtained by 0.2 µm filtration from diatom cultures, with or without nutrient reduction, and from natural diatom populations. The relationship between carbon and TEP was significant, linear and species-specific. Carbon concentration of TEP derived from this relationship concurred with previous findings. Shortage of silicic acid or nitrate in the culture media had no effect on the carbon content of TEP. Molar C:N ratios of TEP were above the Redfield ratio, with a mean value of 26. It is suggested that the nitrogen fraction of TEP can be explained by adsorption of dissolved organic nitrogen (DON) onto TEP. Based on the newly established relationship, concentrations of TEP-derived carbon (TEP-C) were calculated for the Baltic Sea, the coastal Pacific, the North East Atlantic and the Northern Adriatic Sea.

Description: At 2 modern deep-water cold-seep sites, the North Alex Mud Volcano (eastern Mediterranean Sea, water depth ~500 m) and the Concepción Methane Seep Area (south-east Pacific Ocean, water depth ~700 m), we found abundant catshark (Chondrichthyes: Scylio - rhinidae) and skate (Chondrichthyes: Rajidae) egg capsules, respectively, associated with carbonates and tubeworms. Fossilized catshark egg capsules were found at the 35 million year old Bear River Cold-Seep Deposit (Washington State, USA) closely associated with remains of tubeworms and sponges. We suggest that cold-seep ecosystems have served as nurseries for predatory elasmobranch fishes since at least late Eocene time and therewith possibly play an important role for the functioning of deep-water ecosystems.

Description: In 1956, the shallow-water grouper Cephalopholis argus was introduced from Moorea (French Polynesia), where grouper diversity (14 species) is high, to the Main Hawaiian Islands (MHI), where only 2 rare native deep-water groupers occur. In this non-native environment, the species has flourished and has become the dominant apex predator on many reefs. In the present study, a comparison of non-native populations of C. argus in the MHI with native populations in Moorea showed that mean total length (32.0 vs. 26.9 cm), mass (722 vs. 326 g), growth, and body condition were each significantly elevated in the MHI. In addition, while an ontogenetic shift towards larger prey occurred in both locations, it was faster and more consistent in Moorea than in the MHI. As a result, while small C. argus of comparable size in the 2 locations consumed similar-sized prey, large C. argus in Moorea consumed significantly longer and deeper-bodied prey than their counterparts in the MHI. This pattern was unrelated to the size distributions of available prey and may thus reflect stronger intra- and interspecific competition for small prey in Moorea. Although ecological release in a broader sense (i.e. a combination of predator release, parasite release, and competitive release) may play a role, the most direct explanation for the observed differences between C. argus in native habitats in Moorea (with many competing grouper species) and non-native habitats in the MHI (few competitors) would be competitive release (here used in the sense of benefits resulting from the reduction of interspecific competition).

Description: In this study, the impact of oceanic data assimilation on ENSO simulations and predictions is investigated. The authors’ main objective is to compare the impact of the assimilation of sea level observations and three-dimensional temperature measurements relative to each other. Three experiments were performed. In a control run the ocean model was forced with observed winds only, and in two assimilation runs three-dimensional temperatures and sea levels were assimilated one by one. The root-mean-square differences between the model solution and observations were computed and heat content anomalies of the upper 275 m compared to each other. Three ensembles of ENSO forecasts were performed additionally to investigate the impact of data assimilation on ENSO predictions. In a control ensemble a hybrid coupled ocean–atmosphere model was initialized with observed winds only, while either three-dimensional temperatures or sea level data were assimilated during the initialization phase in two additional forecast ensembles. The predicted sea surface temperature anomalies were averaged over the eastern equatorial Pacific and compared to observations. Two different objective skill measures were computed to evaluate the impact of data assimilation on ENSO forecasts. The authors’ experiments indicate that sea level observations contain useful information and that this information can be inserted successfully into an oceanic general circulation model. It is inferred from the forecast ensembles that the benefit of sea level and temperature assimilation is comparable. However, the positive impact of sea level assimilation could be shown more clearly when the forecasted temperature differences rather than the temperature anomalies themselves were compared with observations.

Description: We analysed carbon (δ13C) and nitrogen (δ15N) isotope ratios of organisms and biogenic tissues from Comau Fjord (southern Chile) to characterise benthic food webs and spatial isotope variability in this ecosystem. These values were intended to serve as a baseline for detecting anthropogenic impacts on Patagonian marine fjord ecosystems in later studies. Benthic macro algae and invertebrate suspension feeders were primarily considered, with some supplementary data from cyanobacteria, plankton, fish, and coastal vertebrates. Six depth transects typified the lateral salinity gradients from the innermost part of the fjord to its mouth, as well as the vertical density gradients caused by freshwater inflow. Carbon isotope signatures indicated predominant consumption of either CO2 or HCO3– for benthic macroalgal. All CO2 users belonged to rhodophytes. The δ15N values of benthic macrophytes decreased with decreasing salinity, both vertically and along the fjord axis. This implies the influence of 15N-poor terrestrial dissolved inorganic nitrogen (DIN) at these sites. Enhanced influence of freshwater influx also lowered N contents and increased C/N ratios in algal tissues. Exceptionally high macroalgae δ15N values at the seabird and sealion colony Isla Liliguapi point to animal faeces as an additional source of 15N-enriched DIN. Thus, DIN sources not originating from the open sea are additionally utilised by the benthic macroalgae in the fjord. In contrast, mussel tissue from the same locations was much less influenced by varying DIN sources. Among benthic suspension feeders, mytilids (Mytilus chilensis, Aulacomya ater) had the lowest and scleractinian corals (Desmophyllum dianthus) had the highest δ15N values, and Balanidae (Elminius kingii) and gorgonians (Primnoella sp.) showed values in between. The preference for specific size classes of marine particulate organic matter (seston) as food serves as an explanation for the δ15N variability observed between the different benthic suspension feeders.

Description: Through the use of mesocosm experiments, we show that an unusually early spring phytoplankton bloom can be induced by intermittent high-light periods. We performed mesocosm experiments where plankton assemblages from Kiel Bight (Western Baltic Sea) received a light regime based on the natural seasonal irradiance dimmed to 43% of surface irradiance of cloudless days, starting with irradiance levels of mid-January (6 mesocosms) and mid-February (6 mesocosms). After 6 d, half of the mesocosms received a ca. 2-fold increase in irradiance. In the January mesocosms, a phytoplankton bloom developed only in the treatments with the high-light episode, whereas in the February mesocosms a phytoplankton bloom also developed in the controls. Phytoplankton net growth rates, production:biomass ratios and biomass at the end of the high irradiance episodes were positively correlated to the daily light dose. The relative biomass of diatoms increased with increasing light, whereas the relative biomass of cryptophytes decreased. A bottom-up transmission to mesozooplankton (mainly copepods of the genera Acartia and Oithona) was evident by increased densities of copepod nauplii and egg production under higher light conditions, whereas copepodids and adults showed no responses during the experimental period. The taxonomic composition of the nauplii was shifted to the advantage of Acartia/Centropages (not distinguished at the naupliar stage) under higher light conditions.

Description: The Redfield N:C ratio is a fundamental quantity in marine biogeochemistry because it is a key determinant of the efficiency of the biological carbon pump, yet no convincing explanations have been put forward for its remarkable constancy over much of the world ocean. Phytoplankton growth models have so far been unable to account for the different relationships between growth rate and N:C ratio under nutrient and light limitation, and have not been able to predict the Redfield N:C ratio. A relatively simple model of coupled chlorophyll and nutrient dynamics is developed from the premise that phytoplankton maximize growth by optimally allocating nutrient and energy resources among com- peting metabolic requirements for nutrient uptake, light-harvesting, and growth. The model reconciles nutrient and light limitation and appears valid under both balanced and non-balanced growth condi- tions. The Redfield N:C ratio and its constancy are explained as a result of evolutionary pressure towards maximizing light-limited growth rates in relatively carbon-rich oceanic waters.

Description: In an effort to identify the key mechanisms controlling biological productivity and food web structure in the Chesapeake Bay estuarine turbidity maxima (ETM), we measured plankton community metabolism on a series of surveys in the upper Chesapeake Bay during the winter and spring of 2007 and 2008. Measured quantities included primary production, bacterial production, planktonic community respiration, and algal pigment concentrations. These measurements revealed a classic minimum in photosynthesis in the vicinity of the ETM. Temporal variability in plankton community metabolism, primary production, respiration, and bacterial production, were highest in the southern oligohaline region down-estuary of the ETM, and appeared to be driven by dynamic bio-physical interactions. Elevated primary production and community respiration in this region were often associated with the presence of mixotrophic dinoflagellates. The dinoflagellate contribution to primary production and respiration appeared to be particularly large as a result of their mixotrophic capabilities, which allow them to obtain energy both autotrophically and heterotrophically. This study suggests that mixotrophic dinoflagellates play a key role in pelagic food web in the oligohaline region of Chesapeake Bay supplying most of the labile organic matter during late winter and spring and also providing a vector for transferring microbial production to mesozooplankton.

Description: Diverse coastal seaweed communities dominated by perennial fucoids become replaced by species-poor turfs of annual algae throughout the Baltic Sea. A large-scale field survey and factorial field experiments indicated that grazers maintain the fucoid community through selective consumption of annual algae. Interactive effects between grazers and dormant propagules of annual algae, stored in a 'marine seed bank', determine the response of this system to anthropogenic nutrient loading. Nutrients override grazer control and accelerate the loss of algal diversity in the presence but not in the absence of a propagule bank. This implies a novel role of propagule banks for community regulation and ecosystem response to marine eutrophication.

Description: In the Baltic Sea, rye tested how short nutrient pulses of different lengths and frequencies affect macroalgae, epiphytes, grazers and their interactions. We hypothesized that even small-scale variations in nutrient supply may have significant impacts by favoring fast-growing epiphytes which can cause large-scale declines of canopy-forming macroalgae. In a factorial field experiment single plants of the canopy-forming macroalga Fucus vesiculosus with and without epiphytes were exposed to pulses of elevated nutrients (N and P) over 25 d. Five 1 h pulses given every 5 d had no significant effects. A single 5 h pulse increased the epiphyte load but not F. vesiculosus growth rate. In contrast, increasing epiphyte load caused F. vesiculosus growth rate to decline and attracted higher densities of gastropod grazers. These results indicate that a single nutrient pulse can have rapid direct and indirect effects on macroalgae and their associated epiphytes and grazers. Temporal variability of nutrient supply (five 1 h vs one 5 h pulse) plays a significant role in determining the response of primary producers and consumers to elevated nutrients.

Description: An experimental laboratory set-up was used to study the influence of different grain size compositions and temperatures on the growth of benthic cyanobacteria and diatoms, and on the competition between these 2 groups. Monospecific cultures of 3 species of cyanobacteria (Merismopedia punctata, Microcoleus chthonoplastes, Oscillatoria limosa), and of 2 species of benthic diatoms (Phaeodactylum tricornutum and Nitzschia sp.) were used. The organisms were cultured in 100 ml flasks filled with medium and 3 different kinds of sediment: (1) Sand (fine sand, 63 to 200 µm), (2) Mud-I (mixed fine sand and mud 〈63 µm in the ratio 80:20 wt %), (3) Mud-II (mixed fine sand and mud in the ratio 50:50 wt %). Experimental temperatures were 10, 15 and 25°C. At 10°C and 15°C, both diatom species achieved the highest biomass on the sediments of the finest grain size (50 wt % 〈 63 µm) while cyanobacteria achieved low biomass levels. Coarsening of sediments at the same temperature levels revealed a gradually lower biomass of the diatoms. Particularly on sand, the diatoms never reached the same concentrations of chlorophyll a as on mud. The cyanobacteria, on the other hand, had the highest biomass on sand at 15°C. In the competition experiments the benthic diatom species Nitzschia sp. dominated all types of sediments at 10°C and 15°C. The experiments at 25°C were dominated by the filamentous cyanobacterium M. chthonoplastes. This indicates the importance of abiotic conditions for the distribution and abundance of benthic phototrophic micro-organisms.

Description: The response of epilithic microphytobenthos to in situ nutrient enrichments was studied in the Kiel Fjord, Western Baltic Sea. For this purpose an experimental setup with continuous nutrient supply was designed and installed. Experiments followed the colonization of artificial substrates and the responses of benthic algae to different concentrations and combinations of nitrogen and phosphate. They revealed nitrogen limitation of epilithic microflora from late spring to autumn, such that there was higher biovolume with increasing nitrogen concentrations. Diatoms became dominant in all experiments except one in which the rhodophyte Ceramium strictum prevailed. Species composition was altered by nutrient treatments; one to several species were highly favoured by nutrient enrichment. Consequently, diversity was negatively correlated with final yield. These findings support the hypothesis that competition is an important factor structuring microphytobenthic communities.

Description: Nutrient competition experiments were performed with periphytic microalgae from the Kiel Fjord, western Baltic Sea. Diatoms were dominant competitors at high Si:N ratios (under N-limited conditions) and high Si:P ratios (under P-limited conditions). Under low silicate supply, cyanobacteria became dominant at low N:P ratios and chlorophytes at high ones. Changes in light intensity influenced the outcome of competition at the species level but not at the level of higher taxa. The nitrogen source (ammonium or nitrate) had no impact on the outcome of competition.

Description: The cod Gadus morhua stock in the eastern Baltic Sea has seen a recent miraculous recovery, mainly due to a drastic decrease in fishing mortality (F). Here we explore 3 options for rebuilding the stock to the size (biomass that can produce the maximum sustainable yield, or Bmsy) that is required by international law and that can support high long-term yields. The first option implements a fishing mortality of F = 0.3, as is aimed for under the current European Commission management plan. The second option implements in addition the current constraint of a maximum annual increase of 15% in total allowable catches until a catch corresponding to F = 0.3 is reached. The third option freezes the 2010 catch for 2 yr before allowing a linear increase with spawning stock biomass towards 90% of the maximum sustainable yield. We show that the first option provides the highest catches for the first 3 yr, but fails to rebuild the biomass to the level of Bmsy. The second option rebuilds the biomass above Bmsy only temporarily and results in the lowest catches over a 10 yr period. The third option rebuilds the biomass above Bmsy and provides high catches and the highest profit within 10 yr. Within a decade, all 3 options provide several-fold higher biomasses, catches, and profits compared to the current situation, underlining the benefits that can be obtained from proper fisheries management. In comparison, the so-called precautionary management approach (F = 0.6) implemented until 2006 would perform worse in every respect.

Description: We monitored the dynamics of extracellular organic matter in 3 mesocosms: one dominated by a heterotrophic (microbial) community with negligible autotrophic activity (net heterotrophic system), a second where a small Phaeocystis bloom developed (production and loss almost balanced), and a third harboring a large diatom bloom (net autotrophic system). In all mesocosms, meso- and macroscopic heterotrophic organisms were excluded to primarily study extracellular organic matter production and turnover by specific algae and microbial loop organisms, respectively. Concentration and composition of dissolved organic matter (DOM), i.e. dissolved organic carbon (DOC), monosaccharides and total carbohydrates (MCHO and TCHO), free and combined neutral carbohydrates (DFCHO and DCCHO), as well as free and combined amino acids (DFAA and DCAA) were measured. In addition, net and gross community production rates were determined to calculate C-budgets. Whereas concentrations and composition of MCHO differed very little among the 3 mesocosms, dynamics of TCHO, DFCHO, and DCCHO differed significantly. Concentrations of DFAA were higher in both algal mesocosms compared to the heterotrophic system, and composition of DFAA was significantly different in the Phaeocystis and Diatom tanks. The composition and concentration of DCAA, however, were similar in all 3 mesocosms. Total dissolved carbohydrates and amino acids comprised a substantial fraction of the DOC pool. Dynamics of these DOC fractions, however, could only partly explain those of DOC, implying either that other dissolved compounds were important for overall C-cycling, or that microbial degradation of DOM affects the detection of carbohydrates and protein components.

Description: The effect of phytoplankton on the other compartments of a simple microbial loop consisting of dissolved matter, bacteria and nanoflagellates was investigated in three 1000 l mesocosms. These mesocosms were inoculated with a natural bacterial community and (1) no other additions (No Addition tank), (2) Phaeocystis globosa (Phaeocystis tank), or (3) 2 diatom species (Diatom tank). During the 20 d experiment, autotrophic activity was negligible in the No Addition tank. In contrast a small P. globosa bloom developed in the Phaeocystis tank and a large diatom bloom dominated the Diatom tank. In this paper we describe the experiment, the changes in chlorophyll a and heterotrophic nanoflagellate concentrations, as well as the cycling of nitrogen, phosphorus, and silica. Then we provide a synthesis of the structure and functioning of the microbial loops in these 3 systems using cluster analysis, a statistical pattern recognition tool. The goal was to test the hypothesis that differences in the resident phytoplankton populations would be reflected in (1) the composition and concentration of dissolved organic matter, (2) the composition of the bacterial community, (3) the food web, and (4) the cycling of elements and organic matter. In all 3 mesocosms, nitrate and silicic acid remained abundant. Orthophosphate was preferred by diatoms, whereas Phaeocystis appeared to prefer dissolved organic phosphorus. The hypothesis that phytoplankton composition shapes the structure and functioning of the microbial loop was partially supported: 6 d after inoculation each mesocosm exhibited a distinct organic matter signature. After 10 to 12 d, concentrations of heterotrophic nanoflagellates were high enough to exert significant grazing pressure in all 3 mesocosms. A parallel shift in bacterial community composition was visible in all mesocosms at this time, possibly reflecting grazing pressure. The food-web structure developed divergently in the 3 mesocosms during the second half of the experiment. Differences in biochemical cycling between mesocosms were predominantly driven by the large quantitative differences in autotrophs.

Description: The seasonal cycle over the tropical Pacific simulated by 11 coupled ocean–atmosphere general circulation models (GCMs) is examined. Each model consists of a high-resolution ocean GCM of either the tropical Pacific or near-global means coupled to a moderate- or high-resolution atmospheric GCM, without the use of flux correction. The seasonal behavior of sea surface temperature (SST) and eastern Pacific rainfall is presented for each model. The results show that current state-of-the-art coupled GCMs share important successes and troublesome systematic errors. All 11 models are able to simulate the mean zonal gradient in SST at the equator over the central Pacific. The simulated equatorial cold tongue generally tends to be too strong, too narrow, and extend too far west. SSTs are generally too warm in a broad region west of Peru and in a band near 10°S. This is accompanied in some models by a double intertropical convergence zone (ITCZ) straddling the equator over the eastern Pacific, and in others by an ITCZ that migrates across the equator with the seasons; neither behavior is realistic. There is considerable spread in the simulated seasonal cycles of equatorial SST in the eastern Pacific. Some simulations do capture the annual harmonic quite realistically, although the seasonal cold tongue tends to appear prematurely. Others overestimate the amplitude of the semiannual harmonic. Nonetheless, the results constitute a marked improvement over the simulations of only a few years ago when serious climate drift was still widespread and simulated zonal gradients of SST along the equator were often very weak.

Description: A model, composed of coupled particle tracking and benthic response modules, for predicting waste solids flux and benthic impacts of gilthead sea bream (Sparus aurata L.) and sea bass (Dicentrarchus labrax L.) aquaculture, was tested at six sites with different hydrodynamics, bathymetries and biomasses in the Aegean and Ionian Seas, Eastern Mediterranean with observations of sediment trap flux and benthic impact indicators. Seven sediment trap validation studies were conducted that varied in design with traps deployed either on the sea bed, attached to nets or in the water column. Model predictions of flux to traps spaced 5 m apart up to 50 m from the cages over a 13 d period were statistically significant (r2 = 0.61, n = 57, p ≤ 0.05). However, the model could not predict adequately the flux to traps spaced 2 m apart in the high-flux zone underneath cages where variability between trap observations was high. In this high-flux zone underneath cages, the averaged model flux predictions resulted in a performance of ± 49%. Statistically significant relationships were established at four sites; between modelled flux and either benthic fauna impact indicator species (S), abundance (A), A/S ratio, Shannon Wiener Index or Biomass Fractionation Index (BFI), (r2 = 0.82, 0.60, 0.57, 0.67 and 0.48, respectively; n = 24, p ≤ 0.05). Two other sites, which did not exhibit an abundance peak in enriched zones, did not fit these relationships. Using relative abundance of taxonomic groups, a modelled flux of 4.1 g m-2 d-1 was a useful boundary; on either side of this boundary, clear trends occurred in pollutant tolerant and intolerant species.

Description: Submarine mud volcanism represents an important. pathway for methane from deeper reservoirs to the surface, where it enters the benthic carbon cycle. To quantify overall methane release from the Captain Arutyunov mud volcano (CAMV) and to assess the contribution of macrobenthic seep organisms to the regulation of the benthic methane flux, we linked water column methane concentrations, seabed methane emission and pore water geochemistry to the spatial distribution of seep biota. Prominent organisms of the CAMV seep biota were 3 different species of frenulate tubeworms. Seabed methane emission ranged from 0.001 to 0.66 mmol m(-2) d(-1). Dense patches of tubeworms were associated with the lowest seabed methane emission. Elevated methane emission was associated with a sporadic distribution of tubeworms and the occurrence of numerous mud clasts. Despite the presence of a large subsurface methane reservoir, the estimated total methane release from CAMV was low (0.006 x 10(6) mol yr(-1)). In addition to direct methane consumption by Siboglinum poseidoni, the tubeworms likely contribute to the retention of methane carbon in the sediment by affecting bacterial communities in the proximity of the tubes. The siboglinids create new meso-scale habitats on the sediment Surface, increasing habitat heterogeneity and introducing niches for bacterial communities.

Description: Silver scabbard fish Lepidopus caudatus (Euphrasen, 1788) (Trichiuridae) from the Great Meteor Seamount (GMS) in the central eastern Atlantic were studied for diet composition and metazoan parasites. A total of 36 specimens with lengths between 39.1 and 52.2 cm were sampled, which had taken 14 different prey items belonging to 4 major taxonomic groups (Chaetognatha, Crustacea, Mollusca and Teleostei). The most abundant prey organisms were Myctophidae and Euphausiacea, followed by Copepoda (Calanoida), Decapoda, Chaetognatha and Cephalopoda. Fishes were also the dominant prey in terms of biomass. Cannibalism was observed in 7 specimens of subadult L. caudatus. A total of 11 parasite species were identified in/on L. caudatus. We established 9 new host and 8 new locality records. Infestation rates were congruent with diet composition, indicating that parasites were ingested via mesopelagic prey organisms serving as intermediate hosts. The rich parasite fauna in L. caudatus reflects a high diversity of mesopelagic species at the GMS, providing niches for parasites and their intermediate hosts. While several species such as Paradiplectanotrema lepidopi (Monogenea) and Nybelinia lingualis (Cestoda) are typical parasites of L. caudatus, other species such as Sphyriocephalus tergestinus (Cestoda), Anisakis simplex (Nematoda) and Bolbosoma vasculosum (Acanthocephala) seem to be transferred by hosts migrating into the area, indicating an important role of the GMS in the transoceanic distribution patterns of such parasites

Description: Photoacclimation models are a prerequisite for accurate estimates of primary production in aquatic environments under typically variable light conditions. They generally start from empirical functions of the internal chlorophyll a (chl a) or nutrient quota (e.g. the Droop model). We propose that physiological variations in phytoplankton reflect phenotypic adaptation which maximizes the growth rate. Growth maximization has to account for indirect effects of the enhancement of carbon (C) acquisition by acclimation, primarily through concomitant changes in the intracellular nitrogen (N) budget. Our model expresses, for the first time, the indirect effect of alterations in N uptake on C assimilation by a parameter-free trade-off between the 2 uptake functions. The model explicitly prescribes optimal protein partitioning between N and C uptake and sub-partitioning into carboxylation (1,5-bisphosphate carboxylase/oxygenase, Rubisco) and light harvesting. Applications to various published experimental data for different phytoplankton species support the validity of the optimality hypothesis and point to different flexibility in the re-organization of chloroplasts between taxa as well as to different time-scales on which photoacclimation operates. Simulations of a batch culture with the haptophyte Isochrysis galbana show that a decoupling in pigment N:C from cellular N:C may explain observed lag phases in chl a:C regulation. For diatoms, seemingly stronger constraints in intra-cellular stoichiometry determine the photoacclimative response to variable light regimes, as simulated and reported for Skeletonema costatum. N and chl a quotas correlate well in nutrient-limited chemostats of Thalassiosira fluviatilis, but in part decouple under light limitation. In N limited growth, non-linearity in N:C as expressed by the Droop function results from a combination of a linear quota dependency, down-regulation of relative carboxylation capacity, and increasing N costs of chl a synthesis at elevated growth rates. Our optimality assumption that includes indirect feed-backs through the concept of protein partitioning generates an accurate model for adaptation in physiological traits.

Description: Zooplankton feeding formulations in plankton models have exclusively focused on the relation between food concentration and ingestion, with respiration and excretion being treated separately, despite experimental evidence for strong links among these processes. We present an optimal current-feeding model linking ingestion, respiration, and assimilation efficiency to foraging activity. The Ivlev model is a special case of our optimal current-feeding model, which applies to static feeding behaviour. We validate our model with experimental data for copepods, ciliates, and dinoflagellates. Parameter estimates suggest that phylogenetic grouping is more important than predator size in determining feeding behaviour. Respiratory costs of foraging, e.g. for generating a feeding current, may be much larger than previously thought, are larger in smaller organisms, and might explain the independent development of feeding thresholds in different micro- and mesozooplankton groups. Both preferential feeding on, and lower feeding thresholds for, larger food particles are predicted to derive from greater capture efficiency owing to enhanced detectability of larger particles. The relation between feeding threshold and prey size appears to depend on feeding strategy but not on predator size, as a common relationship seems to apply for current feeders (ciliates and copepods) spanning a vast size range. Our model exhibits an inverse relationship between ingestion and assimilation efficiency, reducing the contribution of copepods to export of organic matter relative to remineralisation at low food concentrations. Export ratio variations previously thought to require strong shifts in community composition can be generated by changes in feeding behaviour predicted by our model.

Description: The impact of moderate environmental stress may be modulated by stress-induced shifts of biotic interactions such as host – epibiont relationships. We studied the stress regime in shallow Western Baltic habitats, the variability of fouling at different temporal and spatial scales, and whether common stressors - low light, high temperature, grazing – affect the abundance and composition of the biofilm on a regionally important macroalga, the bladder wrack Fucus vesiculosus. We further explore the alga’s capacity to chemically modulate the recruitment of microfoulers and whether this ability is impacted by stress. In laboratory, mesocosm and field experiments fouling pressure and epibiotic cover on the algae varied strongly with changing environmental conditions such as temperature, irradiance, depth or grazing. The expectation that abiotic stress affects the fouling-modulating ability of the alga and, thus, indirectly produces the observed variability of epibiosis was not generally confirmed. Indeed, while the strength of chemical antifouling resistance varied seasonally, with a maximum in winter/spring and a minimum in late summer, this could not be related to temporal patterns of environmental stress, fouling pressure, or growth of Fucus. Only the seasonal variation in reproduction seemed to be in phase with antifouling activity. Controlled experiments confirmed that resistance strength was not affected by temperature or grazing, and only moderately by light. We conclude that the fouling modulation ability of Fucus vesiculosus may suffer from light-reduction (e.g. by eutrophication effects) while they are not sensitive to the predicted warming or enhanced grazing.