ALBERT

Beschreibung: Growth rates of the cold-water corals (CWC) Madrepora oculata, Lophelia pertusa, Desmophyllum dianthus and Dendrophyllia cornigera were measured over 8 mo under controlled conditions (12°C in the dark, fed 5 times a week) by means of the buoyant weight technique. Additionally, linear growth rates were measured in M. oculata and L. pertusa for 2 and 1 yr, respectively. The weight measurements revealed growth rates, expressed as percent growth per day (mean ± SD), of 0.11 ± 0.04 for M. oculata, 0.02 ± 0.01 for L. pertusa, 0.06 ± 0.03 for D. dianthus and 0.04 ± 0.02 % d–1 for D. cornigera. Growth in M. oculata was significantly higher (p 〈 0.0001) than in the other 3 CWC species. For M. oculata and L. pertusa, also linear growth was recorded. These values (mean ± SD) were 0.014 ± 0.007 and 0.024 ± 0.018 mm d–1 for M. oculata and L. pertusa, respectively. This is the first study that compares the growth rates of 4 different CWC species under the same experimental conditions of water flow, temperature, salinity and food supply. These corals have different growth rates, both in terms of total weight increase and linear increase, and these growth rates can be related to interspecific physiological differences. Data on growth rates are essential to understand the population dynamics of CWC as well as the recovery capacity of these communities after disturbance.

Beschreibung: Mucus, a complex composed primarily of carbohydrates, is released in similar quantities by scleractinian warm- and cold-water reef corals, and can function as an important carrier of organic material from corals to a range of consumers, microbes in particular. However, information about mucus chemical composition is rare for warm-water corals and non-existent for cold-water corals. This study therefore presents comparative carbohydrate composition analyses of mucus released by the dominant and cosmopolitan warm- and cold-water coral genera. Arabinose was the major mucus carbohydrate component for the genus Acropora, but was not found in cold-water coral mucus. Mucus derived from corals of the genus Fungia contained significantly more fucose than the mucus of all other coral genera. However, comparison of mucus carbohydrate composition for the warm- and cold-water corals in the present study and in the literature revealed no significant differences. This indicates use of similar carbohydrate components (with the exception of arabinose) during mucus synthesis by scleractinian corals, largely irrespective of zooxanthellate or azooxanthellate carbon supply mechanisms.

Beschreibung: The only known population of coelacanths, in the Comores, western Indian Ocean, is endangered by human predation. Historical catch data from Grande Comore reveal that annual catch rates increased steadily from 1954 until the 1970s. This trend was temporarily interrupted due to an international policy introducing motorized boats and promoting offshore fishing techniques. Coelacanths are only caught from traditional unmotorized outrigger canoes as an incidental by-catch of deep water line fishing. A complete survey of all motorized and unmotorized vessels in 1995 at Grande Comore in comparison to earlier years indicated that a recent decreased use of motors and increase of unmotorized canoe fishing has led to an increase in coelacanth catches. Conservation measures and strategies for reducing the fishing pressure exerted on coelacanths are discussed. The southwest coast of Grande Comore should be designated as a nature reserve and protected area where immediate protection measures should be taken, an opinion which is supported by Comorian authorities.

Beschreibung: VHF telemetry was used in November and December 1995 on 8 Humboldt penguins Spheniscus humboldti breeding at Pan de Azúcar Island (26°S, 70°W), Northern Chile, to determine at-sea behaviour of the birds. We obtained 2710 locations, 90% of which were within a radius of 20 km around the island. Mean travelling speed of the birds was 0.92 m s-1 and speed distribution showed peaks at 1.6 and 3 m s-1. Penguins travelling between foraging areas remained submerged for an average of 8.4 s between surfacings, whereas foraging dives lasted on average 61 s. The analysis of 79 complete foraging trips showed that tracks deviated from a straight course, and range (maximum distance from island) was only 0.37 times total horizontal distance swum. Birds did not forage synchronously or in the same foraging areas. However, foraging ranges were correlated between birds, indicating similar search strategies during periods of low food availability. The results obtained here via VHF telemetry agreed well with those of previous studies employing satellite transmitters and data loggers.

Beschreibung: The tissue of 31 demosponge and 7 hexachnelhd species was analyzed for its composition of organic and lnorganic matter With one excephon (Hahclona cf gausaana) inorganic matter i e n~ostly the siliceous skeleton, accounted for most of the dry weight, varying between about 60 and 95%dry wt There were no general trends in the ratio of organic to inorganic matter within sponge orders or genera, and within one species, the ratio could vary between stations For one of the hexactinellids Bathydorus spmosus, several size classes were analyzed and there was no systematic change in the organic inorganic matter ratio w t h specimen size For some species the results from the Weddell Sea sponges are in good agreement with earlier data from McMurdo Sound The low organic matter content in Weddell Sea sponges mphes that sponge biomasses are much lower than hitherto assumed on the basis of then high abundances and large sizes In consequence Antarctic sponges despite their ubiquitousness, may only channel a m n o r fractlon of the general bentho-pelagic flow of matter and energy and their maln role in the ecosystem is likely structural rather than dynamic

Beschreibung: During 5 consecutive summer seasons (1990 to 1994) both Aureliaaurita medusa and mesozooplankton abundances were investigated at 2 stations in the Kiel Bight (western Baltic Sea). Stocks of medusae varied considerably between the years, with median abundance of 1 and 0.3 ind. 100 m-3 in 1990 and 1991, 3 and 4 ind. 100 m-3 in 1992 and 1994, but 9 ind. 100 m-3 in 1993. Significant differences in the mesozooplankton stock and community composition were observed in 1993 when compared with the year of rather low Aurelia abundance (1991). Total zooplankton and copepod numbers both exhibited an inverse relationship with the abundance of medusae and were thus considerably lower in 1993 than in 1991. However, not all copepod species were affected by A. aurita. Pseudo- and Paracalanus spp. and Oithonasimilis showed dramatically reduced stocks in the bloom year when compared with the medusa-poor situation, but no significant changes were found for Centropageshamatus and Acartia spp. Also other zooplankton groups with the exception of bivalve larvae were reduced by the medusae. The differential response of zooplankton to varying abundance of medusae led to a shift in the trophic structure of the zooplankton community. Fine-filter feeders and raptorial feeders were much more important in years when medusae occurred in low densities, whereas coarse-filter feeders dominated in the opposite situation.

Beschreibung: From the Gulf of Aden along a transect to the central-northern Red Sea the abundance and taxonomic composition of metazoan plankton was studied during the southwest monsoon period (summer 1987). Samples were taken with 0.055 mm mesh nets down to a maximum depth of 1050 m. In the epipelagic zone, a distinct decrease in total plankton abundance was observed from south to north, which was much more pronounced in biomass (by a factor of up to 10) as compared to numbers (by a factor of 2). This could partly be explained by differences in the taxonomic and/or size composition of the planktonic fauna. Among non-calanoid copepods, 40 out of 75 species or taxa investigated decreased in abundance from south to north. Sixteen of these species were completely absent in the central-northern area. Nineteen species or taxa, however, showed the opposite feature of a higher abundance in the central-northern Red Sea. The stations were grouped according to similarities in the taxonomic composition of non-calanoid copepods in the epipelagic zone. The following 3 geographical regions could be separated: (1) Gulf of Aden and Strait of Bab al Mandab; (2) southern Red Sea; and (3) central-northern Red Sea. In the meso- and bathypelagic zones, regional differences were not evident. The results are discussed in relation to hydrographic conditions during summer 1987.

Beschreibung: A spring investigation of the phytoplankton in the western Alboran Sea (Mediterranean) was undertaken using chlorophyll and carotenoid biomarkers to characterize the community in the water column and in drifting sediment traps set at 100 and 200 m. During 2 drifter experiments, calm and sunny conditions induced a progressive thermal stratification that reduced pigment sedimentation into deeper water and confined the phytoplankton to the surface layer, resulting in an increase in chlorophyll biomass. 19'-Hexanoyloxyfucoxanthin (prymnesiophytes) and chlorophyll b (chlorophytes, prasinophytes, prochlorophytes) were the major accessory pigments, while fucoxanthin, alloxanthin and peridinin indicated the presence of diatoms, cryptophytes and dinoflagellates, respectively. The proportional contribution of each algal group to the chlorophyll a (chl a) biomass, as derived from multiple regression analysis, revealed that prymnesiophytes, cryptophytes and the green algal group collectively accounted for at least 75% in the upper 100 m, emphasizing the importance of the nanophytoplankton. Phaeopigments, dominated by phaeophorbide a2, were the main pigments observed in sediment traps, although chl a, fucoxanthin and 19'-hexanoyloxyfucoxanthin were detected in smaller concentrations as well as traces of chlorophyll b (chl b). In deep water, fucoxanthin and 19'-hexanoyloxyfucoxanthin were the only accessory pigments present while total phaeopigment/chl a molar ratios 〉1 reflected the active transformation of fine phytogenic material at depth. High particulate organic carbon (POC)/chl a ratios (〉100 in surface water; 〉1000 in deep water) suggested that phytoplankton was a relatively small component of the total carbon biomass down the water column. Using simple budget calculations, we determined that 58 to 65% of the chl a produced in the upper 100 m accumulated in the water column over both experiments. During Expt 1, 29% of the chl a sedimented out, mostly as phaeopigment, at 100 m (24%), and 6% was degraded to colourless residues in the water column. In contrast, only 12% of the chl a sedimented in Expt 2, while 20% was degraded to colourless residues.

Beschreibung: Five hormone-treated female Japanese silver eels Anguilla japonica were tagged with ultrasonic transmitters and released by submersible in the West Pacific at seamounts of the West Mariana Ridge, their supposed spawning grounds. Four eels were tracked for 60 to 423 min in the vicinity of the seamounts. They did not settle at the seamounts but swam at a mean speed of 0.37 m s-1 into open water above deep ground. Their mean swimming depth ranged from 81 to 172 m. Experiments suggest that pre-matured A. japonica migrate to their spawning grounds in temperate warm water and at shallow depths.

Beschreibung: Previous studies have suggested that phytoplankton play an important role in the biogeochemical cycling of iodine, due to the appearance of iodide in the euphotic zone. Changes in the speciation of iodine over the course of the growth cycle were examined in culture media for a variety of phytoplankton taxa (diatoms, dinoflagellates and prymnesiophytes). All species tested showed the apparent ability to reduce iodate to iodide, though production rates varied considerably between species (0.01 to 0.26 nmol l–1 µg–1 chl a d–1), with Eucampia antarctica the least and Pseudo-nitzschia turgiduloides the most efficient iodide producers. Production was found to be species specific and was not related to biomass (indicated by e.g. cell size, cell volume, or chl a content). In all species, except for the mixotrophic dinoflagellate Scrippsiella trochoidea, iodide production commenced in the stationary growth phase and peaked in the senescent phase of the algae, indicating that iodide production is connected to cell senescence. This suggests that iodate reduction results from increased cell permeability, which we hypothesize is due to subsequent reactions of iodate with reduced sulphur species exuded from the cell. A shift from senescence back to the exponential growth phase resulted in a decline in iodide and indicated that phytoplankton-mediated oxidation of iodide to iodate was likely to be occurring. Iodide production could not be observed in healthy cells kept in the dark for short periods. Bacterial processes appeared to play only a minor role in the reduction of iodate to iodide.

Beschreibung: Sensitivity of marine crustaceans to anthropogenic CO2 emissions and the associated acidification of the oceans may be less than that of other, especially lower, invertebrates. However, effects on critical transition phases or carry-over effects between life stages have not comprehensively been explored. Here we report the impact of elevated seawater PCO2 values (3100 µatm) on Hyas araneus during the last 2 weeks of their embryonic development (pre-hatching phase) and during development while in the consecutive zoea I and zoea II larval stages (post-hatching phase). We measured oxygen consumption, dry weight, developmental time and mortality in zoea I to assess changes in performance. Feeding rates and survival under starvation were investigated at different temperatures to detect differences in thermal sensitivities of zoea I and zoea II larvae depending on pre-hatch history. When embryos were pre-exposed to elevated PCO2 during maternal care, mortality increased about 60% under continued CO2 exposure during the zoea I phase. The larvae that moulted into zoea II, displayed a developmental delay by about 20 days compared to larvae exposed to control PCO2 during embryonic and zoeal phases. Elevated PCO2 caused a reduction in zoea I dry weight and feeding rates, while survival of the starved larvae was not affected by the seawater CO2 concentration. In conclusion, CO2 effects on egg masses under maternal care carried over to the first larval stages of crustaceans and reduced their survival and development to levels below those previously reported in studies exclusively focussing on acute PCO2 effects on the larval stages.

Beschreibung: In complex ecosystem models, relationships between species include a large number of direct interactions and indirect effects. In order to unveil some simple and better understandable relationships, it is useful to study the asymmetry of inter-specific effects. We present a simple approach for this based on stochastic food web simulations from previous studies. We refer to the Prince William Sound (Gulf of Alaska) marine ecosystem model for illustration. Real data were used to parameterize a dynamical food web model. Through simulations and sensitivity analysis, we determined the strength of the effects between all species. We calculated the asymmetry between the mutual effects species have on each other, and selected the top 5% most asymmetrical interactions. The set of these highly asymmetrical relationships is illustrated by a separate graph in which we calculated the positional importance of the species and correlated this to other independent properties such as population size and trophic position. Results suggest that halibut is the key species dominating this system of asymmetrical interactions, but sablefish and adult arrowtooth flounder also seem to be of high importance. Nearshore demersals display the highest number of connections in the graph of asymmetrical links, suggesting that this trophic group regulates the dynamics of many species in the food web. This approach identifies key interactions and most asymmetrical relationships, potentially increasing the efficiency of management efforts and aiding conservation efforts.

Beschreibung: The stable carbon isotope composition (δ13C) of particulate organic carbon (POC) was measured in 3 size fractions (POCtotal, POC〉20 µm, POC〈20 µm) during a phytoplankton spring bloom dominated by the diatom Skeletonema costatum in Lindåspollene, a land-locked fjord in southern Norway. In addition to standard parameters for characterizing the phytoplankton bloom (chlorophyll, nutrient, and POC concentrations, and species composition), simultaneous measurements of δ13C of dissolved inorganic carbon (DIC), total alkalinity and DIC concentration were obtained to determine temporal trends in dissolved carbon dioxide concentration and in carbon isotope fractionation (ε p) of the POC size fractions. The carbon isotope composition of the 〉20 µm size fraction, which was dominated by diatoms, was ca 2o/oo heavier than that of the 〈20 µm fraction, which was mainly composed of flagellates. δ13C of both size fractions increased by about 3o/oo over the course of the bloom. A 5o/oo increase in δ13C-PO Ctotal during the bloom resulted partly from a shift in the phytoplankton community from a flagellate- to a diatom-dominated one. Carbon isotope fractionation of all fractions decreased with declining CO2(aq) concentration (14 to 〉6 µmol l-1). A positive correlation between ε p and [CO2(aq)] in the diatom size fraction was obtained for the period of exponential growth. Deviation from this correlation occurred after the peak in cell density and chlorphyll a (chl a) concentration, when POC still continued to increase, and may be related to changing phytoplankton growth rates or to possible effects of nutrient (nitrate) limitation on ε p. Comparison of these results with those of previous field studies shows that, while an inverse relationship is consistently observed between ε p and the ratio of instantaneous growth rate and CO2 concentration {µi/[CO2(aq)]}, considerable scatter exists in this relationship. While this scatter may have partly resulted from inconsistencies between the different studies in estimating phytoplankton growth rate, it could also reflect that factors other than growth rate and CO2 concentration significantly contribute to determining isotope fractionation by marine phytoplankton in the natural environment

Beschreibung: Phosphatase (P-ase) activity was determined together with other extracellular enzyme activities, bacterial abundance and production rates during the 2 SW Monsoon process studies of the German JGOFS Arabian Sea Program. Water samples were collected along the cruise tracks from the equator to the upwelling region at the shelf edge off Oman. Depth profiles of P-ase activity were strikingly different from those of the other enzymes. While values of aminopeptidase and β-glucosidase generally decreased below the euphotic zone, P-ase increased by factors of 1 to 7. The relation between peptidase- and P-ase activity was from 4 to 21 at the surface and from 3 to 5 at 800 m depth. Because P-ase production (dissolved and cell-bound) in deep waters is mainly dependent on bacteria, P-ase activities per bacterial cell were calculated: these were, on average, 37 times higher at 800 m than at the surface. We also observed a positive correlation of P-ase activity with phosphate concentrations in the depth profiles below the euphotic zone, while this relationship was much more variable in the mixed surface layer. These observations suggest that C-limited bacteria in the deep strata did not primarily focus on the phosphate generated by their P-ase activity but on the organic C compounds which were simultaneously produced and which could probably not be taken up prior to the hydrolytic detachment of phosphate. It is hypothesised that a considerable part of the measured P-ase activity was dissolved (though it might have originated from bacteria). These enzymes may be important for the slow, but steady regeneration of phosphate and organic C in mesopelagic waters.

Beschreibung: A predictive model of carbon isotope fractionation (sigma p) and abundance (delta13C phyto) is presented under circumstances where photosynthesis is strictly based on CO2(aq) that passively diffuses into marine phytoplankton cells. Similar to other recent models, the one presented here is based on a formulation where the expression of intracellular enzymatic isotope fractionation relative to that imposed by CO2(aq) transport is scaled by the ratio of intracellular to external [CO2(aq)], ci/ce. Unlike previous models, an explicit calculation of ci is made that is dependent on ce as well as cell radius, cell growth rate, cell membrane permeability to CO2(aq), temperature, and, to a limited extent, pH and salinity. This allows direct scaling of ci/ce to each of these factors, and thus a direct prediction of sigma p and delta13C phyto responses to changes in each of these variables. These responses are described, and, where possible, compared to recent experimental and previous modeling results.

Beschreibung: Stomach contents of 17 sperm whales Physeter macrocephalus stranded in Scotland and Denmark during 1990-96 were analysed. All were sub-adult or adult males and stranded between November and March. They had presumably entered the North Sea during their southward migration from feeding grounds in Arctic waters. Other studies indicate that the majority of the whales were apparently healthy. The diet of these whales was found to consist almost entirely of cephalopods, principally squid of the genus Gonatus (hereafter 'Gonatus', but probably G. fabricii, an oceanic species characteristic of Arctic waters). The other prey species identified were also mostly oceanic cephalopods: the squids Histioteuthis bonnellii, Teuthowenia megalops and Todarodes sagittatus and the octopus Haliphron atlanticus. Although these results are consistent with other recent studies in the area based on single stranded whales, they differ from results of work on whales caught during commercial whaling operations in Icelandic waters (1960s to 1980s) in that little evidence of predation on fish was found in the present study. Remains of single individuals of the veined squid Loligo forbesi, the northern octopus Eledone cirrhosa and the saithe Pollachius virens provided the only possible evidence of feeding in the North Sea. We infer that sperm whales do not enter the North Sea to feed. The timing, and large and uniform sizes of the Gonatus species eaten (most had mantle lengths in the range 195 to 245 mm), as estimated from measurements of the lower beaks, and the seasonality of the strandings is consistent with the whales having fed on mature squid, possibly spawning concentrations--as has recently been reported for bottlenose whales. Assuming that the diet recorded in this study was representative of sperm whales during the feeding season, as much as 500000 t of Gonatus could be removed by sperm whales in Norwegian waters each year and up to 3 times that figure from the eastern North Atlantic as a whole. Evidence from other studies indicates that Gonatus is an important food resource for a wide range of marine predators in Arctic waters.

Beschreibung: Interpretation of diving profiles of aquatic animals would be considerably enhanced by additional behavioural information. A new sensor is presented here which records animal movements. This sensor was tested on a captive loggerhead turtle Caretta caretta which showed similar activity patterns to free-living green turtles Chelonia mydas. A computer program with user-selectable options was developed to analyse the data consistently and rapidly. Using our sensor we calculated the total resting time, which differed by less than 5% from the real resting time when the sampling interval was 2 s. The method was additionally tested for different sampling intervals to find out its applicability for field studies.

Beschreibung: Traditionally, consumer–prey interactions have been considered as purely negative, but herbivores may have positive effects on plants and their productivity. Grazing may enhance prey biomass-specific productivity by directly or indirectly reducing the competition for light, nutrients, and space. We studied the effect of 4 common mesograzers, the isopod Idotea baltica, the amphipod Gammarus oceanicus, and the gastropods Littorina littorea and Rissoa membranacea on epiphytes in an eelgrass Zostera marina L. system. Eelgrass was grown in laboratory mesocosms for a set of experiments manipulating mesograzer species identity, mesograzer density and nutrient concentration. We measured epiphyte biomass-specific productivity via incorporation of radioactive carbon. Herbivore effects on epiphyte photosynthetic capacity were strongly positive for R. membranacea, moderately positive for L. littorea and I. baltica and zero for G. oceanicus under low nutrient supply. Both gastropods increased the nitrogen content of epiphytes, especially the small R. membranacea, and enhanced epiphyte growth. The crustacean species did not increase epiphyte nutrient content, but I. baltica probably enhanced epiphyte productivity by removing the overstory of algal cells, and thus reducing competition for light, nutrients, and space. The positive effect of the 2 gastropod species disappeared under higher nutrient supply, implying the importance of nutrient limitation for this interaction. The positive effect of I. baltica remained at moderate grazer densities despite the higher nutrient concentrations.

Beschreibung: Previous studies on trait-mediated trophic interactions in marine ecosystems were restricted to pair-wise interactions between one species of meso-herbivore and plant, though multi-grazer interactions are more common in nature. We investigated whether the feeding of one consumer, either the periwinkle Littorina littorea or the isopod Idotea baltica, affected consumption by the other consumer via anti-herbivory defence induction in the brown seaweed Fucus vesiculosus. To test the generality of our findings, we ran similar experiments with seaweed/grazer populations in the North and Baltic Seas (NE Atlantic). Grazer-specificity in induction strength was assessed by using the same species of grazer for induction and consumption. ‘Indirect’ induction effects were assessed by using different species of grazers for induction and consumption. Palatability assays were run with live algae and with reconstituted food to distinguish between different mechanisms of resistance. Grazing by herbivores induced a chemical defence in F. vesiculosus. In the North Sea population, the induced defences were only effective against I. baltica, regardless of inducer identity. The sensitive responses of I. baltica to the induced defences were also detected in the reconstituted food assays using Baltic Sea organisms. Thus, marine meso-grazers may be affected by previous feeding through the same or a different species of consumer by modified prey traits, such as induced chemical defences. Furthermore, the magnitude of the effect in the induced defences can be determined by species-specific sensitivity.

Beschreibung: How multiple stressors influence fish stock dynamics is a crucial question in ecology in general and in fisheries science in particular. Using time-series covering a 30 yr period, we show that the body growth of the central Baltic Sea herring Clupea harengus, both in terms of condition and weight-at-age (WAA), has shifted from being mainly driven by hydro-climatic forces to an inter-specific density-dependent control. The shift in the mechanisms of regulation of herring growth is triggered by the abundance of sprat, the main food competitor for herring. Abundances of sprat above the threshold of ~18 × 1010 ind. decouple herring growth from hydro-climatic factors (i.e. salinity), and become the main driver of herring growth variations. At high sprat densities, herring growth is considerably lower than at low sprat levels, regardless of the salinity conditions, indicative of hysteresis in the response of herring growth to salinity changes. The threshold dynamic accurately explains the changes in herring growth during the past 3 decades and in turn contributes to elucidate the parallel drastic drop in herring spawning stock biomass. Studying the interplay between different stressors can provide fundamental information for the management of exploited resources. The management of the central Baltic herring stock should be adaptive and take into consideration the dual response of herring growth to hydro-climatic forces and food-web structure for a sound ecosystem approach to fisheries.

Beschreibung: A total of 354 adult European smelts Osmerus eperlanus (L.) were tested for their ability to survive the screen system of the cooling water inflow of a power plant. With increasing number of musculature parasitic third-stage larvae of Pseudoterranova decipiens, the survival rate of O. eperlanus decreased while the total number of externally visible injuries as well as the number of seriously injured specimens increased. The results indicate that even a single specimen of P. decipiens influences resistance and stamina and affects overall mortality of 7 to 20 cm long smelts. The initial effect of the parasites is to reduce swimming speed of infested fish, which leads to more frequent contact of these fish with the fine meshed screen of the cooling water inlet before they are removed by the automatic cleaning system. If the separated fishes are returned to the main stream, it becomes apparent that the cooling water inflow selectively reduces the number of living parasitised smelt in the area. Thus, the number of parasitic third-stage P. decipiens larvae in the local smelt population which are able to complete their life-cycle is also reduced. P. decipiens makes infested smelt more susceptible to negative anthropogenic influences such as cooling water intake or trawl fisheries.

Beschreibung: Sediment cores of 20 cm diameter contaning the natural benthic fauna were subjected to low oxygen conditions in a laboratory microcosm system. After several days of oxic conditions ('oxic stage') the oxygen content of the water was reduced to 25% saturation for 15 d ('hypoxic stage'), followed by a 'reoxygenation stage'. Effective solute transport rates were calculated using measurements with the conservative tracer ion bromide. Profiles of oxygen and ΣCO2 were measured and molecular diffusive as well as effective fluxes, account mg for effective solute exchange, were calculated. The overall response of the benthic community was to compensate for low oxygen content of the overlying water by increased pumping activity. On average, effective diffusion coefficients (Den} were 3 times higher in hypoxia than under oxic conditions. D eff reached 1.5 x 10^-4 cm2 s^-1, a value 30 times that of molecular diffusion. During hypoxia we observed low molecular diffusive O2 flux, higher effective O2 flux, as well as an increase in ΣCO2 within the sediment. We interpret this as a shift of transport away from diffusion within the bulk sediment interstices (oxic conditions) to the advective transport pathways along burrows during hypoxia. This facilitates fast transport of oxygen and bromide along burrows and contrasts with the slower transport of CO2 from the interstices governed by molecular diffusion. In this transient situation calulations based on gradients result in an unrealistic molar ratio of fluxes(CO2/O2)as high as 11.

Beschreibung: Biological dinitrogen (N2) fixation is the primary input of fixed nitrogen (N) into the marine biosphere, making it an essential process contributing to the biological functions of all organisms. Because biologically available N often limits marine productivity, microbial processes leading to its loss and gain (e.g. denitrification and N2 fixation, respectively) play an important role in global biogeochemical cycles. Bioturbation is known to influence benthic N cycling, most often reported as enhancement of denitrification and a subsequent loss of N2 from the system. N2 fixation has rarely been addressed in bioturbation studies. Instead, sedimentary N2 fixation typically has been considered important in relatively rare, localized habitats such as rhizosphere and phototrophic microbial mat environments. However, the potential for N2 fixation in marine sediments may be more widespread. We show here that nitrogenase activity can be very high (up to 5 nmol C2H4 cm–3 h–1) in coastal sediments bioturbated by the ghost shrimp Neotrypaea californiensis and at depths below 5 cm. Integrated subsurface N2-fixation rates were greater than those previously found for un-vegetated estuarine sediments and were comparable to rates from photosynthetic microbial mats and rhizospheres. Inhibition experiments and genetic analysis showed that this activity was mainly linked to sulfate reduction. Sulfate-reducing bacteria (SRB) are widespread and abundant in marine sediments, with many possessing the genetic capacity to fix N2. Our results show that N2 fixation by SRB in bioturbated sediments may be an important process leading to new N input into marine sediments. Given the ubiquity of bioturbation and of SRB in marine sediments, this overlooked benthic N2 fixation may play an important role in marine N and carbon (C) cycles.

Beschreibung: The focus of this study was the 11.55 m long sediment core 303700-7, which was retrieved from the Gdansk Basin during a cruise of RV ‘Poseidon’ within the frame of the Russian–German Project GISEB. The core was analysed for grain size, elemental chemical composition, organic carbon and palynological spectra. The age control was based on palynostratigraphy and 7 radiocarbon datings of bulk sedimentary organic matter. These data provide a high-resolution record of climatic and marine palaeoenvironments in the Gdansk Basin for the last ca. 13 kyr, from the Bølling to late Holocene time. Sedimentation rates were estimated to vary between 0.37 and 1.62 mm yr–1. Major variations in palaeosalinity were estimated from bromine concentrations in the sediment. This method allows the first quantitative reconstruction of palaeosalinity changes in the Baltic Sea, especially profound during the Littorina and Postlittorina periods (middle to late Holocene). In addition, grain size data indicated several Littorina transgressive–regressive stages and a few episodes of increased near-bottom current activity. Our results from the Gdansk Basin are consistent with palaeoceanographic data from other deep basins of the Baltic Sea and provide new insights into the regional Holocene history.

Beschreibung: We report on a novel sponge disease, hereafter termed 'sponge white patch' (SWP), affecting the Caribbean sponge species Amphimedon compressa. SWP is characterized by distinctive white patches of variable size that are found irregularly on the branches of diseased sponges. Nearly 20% of the population of A. compressa at Dry Rocks Reef, Florida, USA, showed symptoms of SWP at the time of investigation (November 2007-July 2010). Approximately 21% of the biomass of SWP individuals was bleached, as determined by volume displacement. Scanning electron microscopy analysis showed severe degradation of bleached tissues. Transmission electron microscopy of the same tissues revealed the presence of a spongin-boring bacterial morphotype that had previously been implicated in sponge disease (Webster et al. 2002; Mar Ecol Prog Ser 232:305-309). This particular morphotype was identified in 8 of 9 diseased A. compressa individuals investigated in this study. A close relative of the aforementioned disease-causing alphaproteobacterium was also isolated from bleached tissues of A. compressa. However, whether the spongin-boring bacteria are true pathogens or merely opportunistic colonizers remains to be investigated. Molecular fingerprinting by denaturing gradient gel electrophoresis (DGGE) demonstrated a distinct shift from the microbiota of healthy A. compressa to a heterogeneous mixture of environmental bacteria, including several phylotypes previously implicated in sponge stress or coral disease. Nevertheless, tissue transplantation experiments conducted in the field failed to demonstrate infectivity from diseased to healthy sponges, leaving the cause of SWP in A. compressa to be identified.

Beschreibung: Nematodes of the family Stilbonematinae are known for their highly specific association with ectosymbiotic bacteria. These worms are members of the meiofauna in marine, sulfide-rich sediments, where they migrate around the redox boundary layer. In this study, bacterial ectosymbionts of 2 species of marine nematodes, Stilbonema sp. and Laxus oneistus, were shown to be capable of the respiratory reduction of nitrate and nitrite (denitrification). The use of these alternative electron acceptors to oxygen by the bacteria allows the animals to migrate into the deeper, anoxic sediments, where they can exploit the sulfide-rich patches of the deeper sediment layers. The accumulation of thiols (sulfide, thiosulfate, sulfate and glutathione) in body tissues of the worms was determined following incubation in the presence of various electron donors (sulfide, thiosulfate) and acceptors (nitrate). In their chemoautotrophic metabolic potential, the ectosymbionts of the 2 nematode species were found to resemble the phylogenetically related, intracellular symbionts of macrofaunal hosts of deep-sea hydrothermal vents and other sulfide-rich habitats.

Beschreibung: We examined the simultaneous effect of climate warming and light availability on the phytoplankton spring bloom using 1400 l (1 m depth) indoor mesocosms. The timing of the spring bloom was advanced both by warming and higher light intensity, but the influence of temperature on the phytoplankton community was stronger than the light effect. Warming affected phytoplankton directly and indirectly via enhanced grazing pressure at higher temperatures. Warming resulted in markedly lower phytoplankton biomass and a shift towards smaller cell sizes. It also led to changes in the community structure of phytoplankton and zooplankton. Among phytoplankton, large-celled diatoms were most negatively affected by warming. Overwintering zooplankton species (Oithona, Pseudocalanus) remained dominant in the cold treatments, while they were replaced by late spring or summer species (Acartia, Centropages, Temora) in the warmed treatments. Our results show that understanding food web interactions might be very important to the study of the effects of climate warming on pelagic ecosystems.

Beschreibung: We assessed the effects of light limitation and temperature shift on palatability and induced antiherbivore defense in the brown alga Fucus vesiculosus L. Incubation for 2 wk at light intensities above the compensation point of photosynthesis and in the absence of grazers increased the palatability of F. vesiculosus and its subsequent consumption by the omnivorous isopod Idotea baltica Pallas. This effect correlated with an increased C:N ratio and mannitol content in the algal tissue, presumably due to increased photosynthetic carbon fixation. Mannitol, the primary product of photosynthesis in F. vesiculosus, proved to be a feeding cue for I. baltica, and depletion of the mannitol pool may therefore account for the reduced palatability during light limitation. At light intensities above the compensation point of photosynthesis, F. vesiculosus responded with decreasing palatability when it was exposed to I. baltica grazing. Irrespective of the preceding light regime, such defense induction was prevented during incubation under light limitation. Thus, under low light, defense induction is not only inhibited, but also less necessary due to the relative absence of feeding cues. Upward or downward shifts in water temperature by approximately 10°C also inhibited inducible defense in F. vesiculosus. However, such shifts did not affect algal growth and were therefore the consequence of an impairment of specific defense-related components rather than of resource limitation, unless compensatory growth was given priority over defense.

Beschreibung: The synergistic effects of fishing, climate and internal dynamics on population fluctuations are poorly understood due to the complexity of these interactions. In this paper, we combine time series analysis and simulations to investigate the long-term dynamics of an overexploited population in the Mediterranean Sea, and its link with both fishing-induced demographic changes and hydroclimatic variability. We show that the cyclicity of the catch per unit of effort (CPUE) of European hake Merluccius merluccius (EH) vanished in the 1980s, while the correlation between the CPUE and a local environmental index increased. Using simulations, we then show that the cyclicity observed in the EH biomass before the 1980s can have an internal origin, while that its disappearance could be due to the fishing-induced erosion of the age structure. Our results suggest that fishing can trigger a switch from internally generated to externally forced population fluctuations, the latter being characterised by an increasing dependency of the population on recruitment and ultimately on environmental variability. Hydroclimatic modifications occurring in the Mediterranean in the early 1980s could have enhanced these changes by leading to a mismatch between early life stages of EH and favorable environmental conditions. Our conclusions underline the key effect of the interaction between exploitation and climate on the dynamics of EH and its important consequences for management and conservation.

Beschreibung: Biological dinitrogen (N2) fixation is the primary input of fixed nitrogen (N) into the marine biosphere, making it an essential process contributing to the biological functions of all organisms. Because biologically available N often limits marine productivity, microbial processes leading to its loss and gain (e.g. denitrification and N2 fixation, respectively) play an important role in global biogeochemical cycles. Bioturbation is known to influence benthic N cycling, most often reported as enhancement of denitrification and a subsequent loss of N2 from the system. N2 fixation has rarely been addressed in bioturbation studies. Instead, sedimentary N2 fixation typically has been considered important in relatively rare, localized habitats such as rhizosphere and phototrophic microbial mat environments. However, the potential for N2 fixation in marine sediments may be more widespread. We show here that nitrogenase activity can be very high (up to 5 nmol C2H4 cm–3 h–1) in coastal sediments bioturbated by the ghost shrimp Neotrypaea californiensis and at depths below 5 cm. Integrated subsurface N2-fixation rates were greater than those previously found for un-vegetated estuarine sediments and were comparable to rates from photosynthetic microbial mats and rhizospheres. Inhibition experiments and genetic analysis showed that this activity was mainly linked to sulfate reduction. Sulfatereducing bacteria (SRB) are widespread and abundant in marine sediments, with many possessing the genetic capacity to fix N2. Our results show that N2 fixation by SRB in bioturbated sediments may be an important process leading to new N input into marine sediments. Given the ubiquity of bioturbation and of SRB in marine sediments, this overlooked benthic N2 fixation may play an important role in marine N and carbon (C) cycles.

Beschreibung: The warming trend for the entire globe (1850 to 2005) is 0.04°C decade–1. A specific warming period started around 1980 and continues until the present. This warming also occurred in the Baltic Sea catchment, which lies between maritime temperate and continental subarctic climate zones. A detailed study of climate variability and the associated impact on the Baltic Sea area for the period 1958 to 2009 revealed that the recent changes in the warming trend are associated with changes in large-scale atmospheric circulation over the North Atlantic. The number and pathways of deep cyclones changed considerably in line with an eastward shift of the North Atlantic Oscillation centers of action. There is a seasonal shift of strong wind events from autumn to winter and early spring. Since the late 1980s, the winter season (DJFM, i.e. December to March) of the Baltic Sea area has tended to be warmer, with less ice coverage and warmer sea surface temperatures, especially pronounced in the northern parts of the Baltic Sea. There is a tendency for increased cloud cover and precipitation in regions that are exposed to westerlies and less cloud coverage at the leeward side of the Scandinavian Mountains and over the Baltic Sea Basin.

Beschreibung: The role of tetrathionate in the sulfur cycle of Baltic Sea sediments was investigated in different habitats and under a variety of environmental conditions. Sediment profiles were recorded with regard to numbers of thiosulfate oxidizing bacteria, concentrations of sulfur compounds, and potential rates of thiosulfate oxidation. Products of thiosulfate oxidation were quantified in incubated sediment samples and in pure cultures. Evidence was found that tetrathionate is formed within these sediments, that sulfur oxidizing bacteria are present in considerable numbers, that these bacteria are of major importance in the oxidation of reduced sulfur compounds in their habitat, and that tetrathionate is an important oxidation product of these bacteria. Thiosulfate is oxidized by bacteria isolated from these sediments to varying proportions of tetrathionate, sulfate, and also elemental sulfur. In highly sulfidic sediments and in the presence of large amounts of organic matter, tetrathionate was present in sediment horizons in which thiosulfate and elemental sulfur also accumulated. A tetrathionate cycle is proposed to be active in natural marine and brackish water sediments in which, due to combined bacterial action and chemical reactions, a net oxidation of sulfide to elemental sulfur occurs in the presence of catalytic amounts of thiosulfate and tetrathionate.

Beschreibung: The effect of variable CO2 concentrations on the elemental composition (C:N:P) of marine diatoms was investigated in 2 strains of Skeletonema costatum (Grev.) Cleve. Five or 6 concentrations of dissolved molecular carbon dioxide [CO2 (aq)], ranging from 0.5 to 39 µmol l-1, were applied in dilute batch cultures. In both strains, elemental ratios were clearly dependent on [CO2 (aq)]. With decreasing CO2 concentrations, a decline in C:P and N:P and an increase in C:N was observed. The close correlation between C:P or N:P and [CO2 (aq)] corresponded to a ca 45 to 65% decrease in elemental ratios from highest (〉=30 µmol l-1) to lowest (ca 1 µmol l-1) CO2 concentrations. C:N at low [CO2 (aq)] was up to 24% higher than at high [CO2 (aq)]. To date, the elemental composition of marine phytoplankton has been considered to be independent of CO2 availability. If dependency of the C:N:P ratio on [CO2 (aq)] proves to be a general phenomenon in marine phytoplankton, changes in the elemental composition may be expected in response to the currently observed increase in partial pressure of atmospheric CO2.

Beschreibung: During the past few decades, the green crab Carcinus maenas, a native to Europe, has invaded the North American Pacific coast. In this new habitat, C. maenas encounters North American periwinkles of the genus Littorina that differ from European Littorina spp. in size, shape and shell strength. We hypothesize that the ability to handle prey never encountered previously is a prerequisite for successful invasion of novel habitats. In a first approach to testing our hypothesis, we compared European (native) to Canadian (invaded) C. maenas in feeding trials with Littorina spp. from Europe as well as Canada. Canadian crabs had significantly larger crusher claws than European crabs of the same size. Prey handling by Baltic crabs, but not by North Sea crabs, significantly depended on shell morphometry and strength of European periwinkles. By contrast, neither European nor Canadian crabs were affected by shell characteristics of the relatively soft-shelled Canadian periwinkles. Baltic and Canadian crabs did not differ in terms of handling time for, and handling success of, different periwinkle species, but North Sea crabs needed more time for, and were less successful in, crushing periwinkles. We conclude that C. maenas exhibits plasticity in both claw morphometry and feeding behaviour that enables this predator to handle novel prey organisms, and contributes to its success as an invader.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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).

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: We review research from the last 40 yr on macroalgal–bacterial interactions. Marine macroalgae have been challenged throughout their evolution by microorganisms and have developed in a world of microbes. Therefore, it is not surprising that a complex array of interactions has evolved between macroalgae and bacteria which basically depends on chemical interactions of various kinds. Bacteria specifically associate with particular macroalgal species and even to certain parts of the algal body. Although the mechanisms of this specificity have not yet been fully elucidated, ecological functions have been demonstrated for some of the associations. Though some of the chemical response mechanisms can be clearly attributed to either the alga or to its epibiont, in many cases the producers as well as the mechanisms triggering the biosynthesis of the biologically active compounds remain ambiguous. Positive macroalgal–bacterial interactions include phytohormone production, morphogenesis of macroalgae triggered by bacterial products, specific antibiotic activities affecting epibionts and elicitation of oxidative burst mechanisms. Some bacteria are able to prevent biofouling or pathogen invasion, or extend the defense mechanisms of the macroalgae itself. Deleterious macroalgal–bacterial interactions induce or generate algal diseases. To inhibit settlement, growth and biofilm formation by bacteria, macroalgae influence bacterial metabolism and quorum sensing, and produce antibiotic compounds. There is a strong need to investigate the bacterial communities living on different coexisting macroalgae using new technologies, but also to investigate the production, localization and secretion of the biological active metabolites involved in those possible ecological interactions.

Beschreibung: The vertical distribution of methane- and ammonia-oxidizing bacteria (MOB and AOB, respectively), and the physicochemical conditions in the chemocline of Lake Kinneret (Israel) were studied at a resolution of 10 cm from 16.2 to 17.7 m depth. Profiles of the chemical parameters indicated decreasing concentrations of methane (from 22.4 to 0.11 µmol l–1) and ammonia (from 14.2 to 8.4 µmol l–1) towards the water surface and in close proximity to the chemocline. The disappearance of methane coincided with methane oxidation that could be corroborated throughout this layer with highest rates at 17.4 to 17.6 m. Disappearance of ammonia could not be linked to ammonia oxidation exclusively. The genes pmoA and the homologous amoA (coding for subunit α of the methane and ammonia monooxygenase, respectively) were amplified by PCR. The products were analyzed by terminal restriction fragment length polymorphism (T-RFLP) and sequencing of clone libraries. The results demonstrated that different MOB and AOB communities are established along the concentration gradient within the narrow layer of the metalimnetic chemocline. Changes in the intensity of the T-RFLP peaks and the frequency of different groups of alpha- and gammaproteobacterial MOB, and betaproteobacterial AOB, coincided with the concentration gradients of methane, ammonia, nitrate, and oxygen in the chemocline. This suggests that different communities of MOB, and to a lesser extent AOB, contribute to the formation of chemical gradients of their particular substrates in the chemocline

Beschreibung: Rising atmospheric CO2 concentrations could cause a calcium carbonate subsaturation of Arctic surface waters in the next 20 yr, making these waters corrosive for calcareous organisms. It is presently unknown what effects this will have on Arctic calcifying organisms and the ecosystems of which they are integral components. So far, acidification effects on crustose coralline red algae (CCA) have only been studied in tropical and Mediterranean species. In this work, we investigated calcification rates of the CCA Lithothamnion glaciale collected in northwest Svalbard in laboratory experiments under future atmospheric CO2 concentrations. The algae were exposed to simulated Arctic summer and winter light conditions in 2 separate experiments at optimum growth temperatures. We found a significant negative effect of increased CO2 levels on the net calcification rates of L. glaciale in both experiments. Annual mean net dissolution of L. glaciale was estimated to start at an aragonite saturation state between 1.1 and 0.9 which is projected to occur in parts of the Arctic surface ocean between 2030 and 2050 if emissions follow ‘business as usual’ scenarios (SRES A2; IPCC 2007). The massive skeleton of CCA, which consist of more than 80% calcium carbonate, is considered crucial to withstanding natural stresses such as water movement, overgrowth or grazing. The observed strong negative response of this Arctic CCA to increased CO2 levels suggests severe threats of the projected ocean acidification for an important habitat provider in the Arctic coastal ocean.

Beschreibung: Both ocean warming and acidification have been demonstrated to affect the growth, performance and reproductive success of calcifying invertebrates. However, relatively little is known regarding how such environmental change may affect interspecific interactions. We separately treated green crabs Carcinus maenas and periwinkles Littorina littorea under conditions that mimicked either ambient conditions (control) or warming and acidification, both separately and in combination, for 5 mo. After 5 mo, the predators, prey and predator-prey interactions were screened for changes in response to environmental change. Acidification negatively affected the closer-muscle length of the crusher chela and correspondingly the claw-strength increment in C. maenas. The effects of warming and/or acidification on L. littorea were less consistent but indicated weaker shells in response to acidification. On the community level, however, we found no evidence that predator-prey interactions will change in the future. Further experiments exploring the impacts of warming and acidification on key ecological interactions are needed instead of basing predictions of ecosystem change solely on species-specific responses to environmental change.

Beschreibung: The multivariate patterns resulting from analyses of macrobenthic abundance data at different taxonomic levels are compared to the pattern derived from various measurements obtained through sediment profiling imagery (SPI). A time-series data set from 1 station in Kiel Bay (Western Baltic) at 22 m depth including macrobenthic and SPI replicates covering 8 yr (1989 to 1996) was analyzed by means of multidimensional scaling (MDS) ordination. The macrobenthos data showed similar patterns, and there was little information loss, with decreasing taxonomic resolution from species to phylum level. The multivariate pattern in the SPI data was not significantly correlated to any of the macrofaunal patterns. However, macrofaunal and SPI patterns seemed to be complementary since they emphasized different aspects of the long-term succession in the Southern Baltic Sea. While macrofaunal patterns were sensitive to anoxia events, changes of SPI-recorded seabed characteristics were primarily related to physical disturbances possibly due to variations in fishing intensity.

Beschreibung: Cod larvae from laboratory rearing experiments aged from 1 to 12 d after hatching, both fed and deprived of food, were analysed. The number of increments on the otolith and the width of these increments were determined together with the RNA/DNA ratios on the same individual larva. Alizarin marking of the otoliths was performed to confirm the formation of daily increments. Cod larvae reared at 6*C formed the first ring right after hatching and deposited increments on a daily basis. A comparison of the measurements between the right and the left lapillus showed that these can differ, if the radius is taken. The lapillae core showed especially high individual variability, whereas the sum of the increments did not differ between both lapilli. Until Day 10 after hatching, while the larvae were still feeding on their yolk, the external food situation did not affect the increment width of the lapilli or the RNA/DNA ratios. In larvae older than 10 d the width of the daily increments was dependent on the nutritional situation and RNA/DNA ratios decreased in starving larvae in comparison to feeding larvae. RNA/DNA ratios and increment widths were correlated.

Beschreibung: Mixed cultures of 4 polar diatoms regularly found in Antarctic pack ice were grown over 20 d in closed bottles at high light (200 to 250 umol photons m-2 s-1) and at 0*C in order to investigate growth physiology and biomass production under conditions simulating the sea ice habitat during summer. Species tested were: Chaetoceros cf. neogracile, Fragilariopsiscylindrus, Thalassiosiraantarctica and Porosira pseudodenticulata. Initially, all species grew exponentially, but exponential growth ceased for P. pseudodenticulata and T.antarctica after 6 d, for F.cylindrus after 8 d, and for C. cf. neogracile after 10 d. Slight increases in cell number were observed for all species 2 d later. Peak biomass amounted to 140 ug chl a (850 umol particulate organic carbon, POC) l-1. At the same time, concentrations of dissolved inorganic carbon (DIC) were reduced by 1000 uM, oxygen concentrations increased to 1400 uM, and pH increased to 10.5. At this stage, a substantial decline in plasma-containing cells was recorded for F.cylindrus. C. cf. neogracile accounted for 80%, and C. cf. neogracile and F.cylindrus accounted for 〉95% of total carbon biomass. The carbon isotope composition of POC (expressed as delta13C) increased from -24 to -9 during the experiment. Model calculations showed that diffusive uptake of dissolved CO2 satisfied cellular carbon demand for all species except P. pseudodenticulata at CO2(aq) concentrations 〉0.5 uM, whereas direct HCO3- utilization was observed for C. cf. neogracile below this concentration. Our data confirm that intense photosynthetic carbon assimilation may lead to profound chemical changes in isolated interstitial brine solutions, with significant consequences for sea ice biota. We propose that the capacity to efficiently utilize ambient DIC, possibly mediated by virtue of favorable surface to volume ratios as well as active pathways of inorganic carbon acquisition, favors growth of small diatoms, and may be an important factor driving ice algal species succession during summer blooms. Since only 2 species continued to grow in fresh medium following experimental incubation (C. cf. neogracile and P. pseudodenticulata), differential tolerance to chemical variations may influence the seeding potential of ice algae following release into the open water.

Beschreibung: Diving seabirds should evolve a variety of foraging characteristics which enable them to minimize energy expenditure and to maximize net energy gain while searching for prey underwater. In order to assess the related ecological adaptations in a marine predator, we studied the at-sea distribution and the diving behaviour of 23 cormorants Phalacrocorax carbo (Linnaeus) breeding at the Chausey Islands (France) using VHF-telemetry and data loggers recording hydrostatic pressure. Birds foraged within an area of approximately 1131 km2 situated north-east of the breeding colony. This zone represents only 25% of the maximal potentially available area that the birds may utilize considering their maximum foraging range of 35 km. Individual birds remained within restricted individual foraging areas (on average 18 and 10% of the total utilized area in 1994 and 1995, respectively) throughout the study period. Moreover, the cormorants studied conducted an average of 42 dives per foraging trip, lasting for an average of 40 s (maximum 152 s), and reached an average maximum dive depth of 6.1 m (maximum 32 m) with median descent and ascent angles calculated to be 18.7° and 20.3°, respectively. Overall, 64% of all dives were U-shaped dives and 36% V-shaped dives. We use these results to demonstrate how both specialization and opportunism may support the remarkably high foraging efficiency of this marine predator.

Beschreibung: Changing seawater chemistry towards reduced pH as a result of increasing atmospheric carbon dioxide (CO2) is affecting oceanic organisms, particularly calcifying species. Responses of non-calcifying consumers are highly variable and mainly mediated through indirect ocean acidification effects induced by changing the biochemical content of their prey, as shown within single species and simple 2-trophic level systems. However, it can be expected that indirect CO2 impacts observed at the single species level are compensated at the ecosystem level by species richness and complex trophic interactions. A dampening of CO2-effects can be further expected for coastal communities adapted to strong natural fluctuations in pCO2, typical for productive coastal habitats. Here we show that a plankton community of the Kiel Fjord was tolerant to CO2 partial pressure (pCO2) levels projected for the end of this century (〈1400 µatm), and only subtle differences were observed at the extremely high value of 4000 µatm. We found similar phyto- and microzooplankton biomass and copepod abundance and egg production across all CO2 treatment levels. Stoichiometric phytoplankton food quality was minimally different at the highest pCO2 treatment, but was far from being potentially limiting for copepods. These results are in contrast to studies that include only a single species, which observe strong indirect CO2 effects for herbivores and suggest limitations of biological responses at the level of organism to community. Although this coastal plankton community was highly tolerant to high fluctuations in pCO2, increase in hypoxia and CO2 uptake by the ocean can aggravate acidification and may lead to pH changes outside the range presently experienced by coastal organisms.

Beschreibung: The notion that excess phosphorus (P) and high irradiance favour pelagic diazotrophy is difficult to reconcile with diazotroph behaviour in laboratory experiments and also with the observed distribution of N2-fixing Trichodesmium, e.g. in the relatively nitrogen (N)-rich North Atlantic Ocean. Nevertheless, this view currently provides the state-of-the-art framework to understand both past dynamics and future evolution of the oceanic fixed N inventory. In an attempt to provide a consistent theoretical underpinning for marine autotrophic N2 fixation we derive controls of diazotrophy from an optimality-based model that accounts for phytoplankton growth and N2 fixation. Our approach differs from existing work in that conditions favourable for diazotrophy are not prescribed but emerge, indirectly, from trade-offs among energy and cellular resource requirements for the acquisition of P, N, and carbon. Our model reproduces laboratory data for a range of ordinary phytoplankton species and Trichodesmium. The model predicts that (1) the optimal strategy for facultative diazotrophy is switching between N2 fixation and using dissolved inorganic nitrogen (DIN) at a threshold DIN concentration; (2) oligotrophy, especially in P and under high light, favours diazotrophy; (3) diazotrophy is compatible with DIN:DIP supply ratios well above Redfield proportions; and (4) communities of diazotrophs competing with ordinary phytoplankton decouple emerging ambient and supply DIN:DIP ratios. Our model predictions appear in line with major observed patterns of diazotrophy in the ocean. The predicted importance of oligotrophy in P extends the present view of N2 fixation beyond a simple control by excess P in the surface ocean.

Beschreibung: Simultaneous analysis of carbon, nitrogen and sulphur stable isotope ratios was applied in this pilot study to examine the food web of a Zostera marina L. system in the western Baltic Sea. Samples of three potential food sources: eelgrass, epiphytic algae and seston, as well as 69 consumer species were collected during the growing season of Z. marina from March to September 2011. The measured δ13C values of epiphytes (-14.1‰ ± 1.8 SD) were close to δ13C values of eelgrass (-11.6‰ ± 1.8 SD), impeding a clear distinction of those two carbon sources, whereas seston δ13C values (-20.9‰ ± 3.5 SD) were clearly different. This frequently encountered problem was solved by the additional use of δ34S, which resulted in easily distinguishable values for sediment and seawater derived sulphur. Values of primary producer δ34S ranged from 5.6‰ (± 2.3 SD) for Z. marina leaves to 14.2‰ (± 1.6 SD) for epiphytes and 11.9‰ (± 3.3 SD) for seston. The combination of δ34S and δ13C values made a separation of carbon sources possible and enabled the allocation of potential food sources to consumers and a description of their trophic relationships. The data of stable isotope ratio analysis of this eelgrass community strongly indicate a food web based on epiphyte and seston production. δ15N values show a food web consisting of large numbers of generalists and a high degree of omnivory amongst the consumer species analysed. This implies an occupation of every trophic position possible, which is supported by a continuous distribution of δ15N values. Previously described eelgrass food webs may have to be re-evaluated to include sulfur in order to provide a clear picture on primary carbon sources.

Beschreibung: Snails Littorina littorea with and without artificial epibionts were exposed in situ for nearly 1 mo. Measurement of individual shell length increase over this period revealed that clean snails grew 3 times faster than fouled snails. These results compare well with previously conducted flume experiments (Wahl 1996). This epibiosis effect is thought to be due to increased drag-the only feature distinguishing the 2 treatment groups-caused by the presence of epibionts on the shell. Increased drag probably entails higher energy expenditure for pedal activities (attachment and locomotion) and a reduced allocation of resources towards growth and, possibly, reproduction

Beschreibung: Simultaneous triple stable isotope analysis of carbon, nitrogen and sulphur was employed to study the temporal variation in the food web of a subtidal eelgrass Zostera marina bed in the western Baltic Sea. Samples of 3 potential food sources (eelgrass, epiphytes and seston) and consumer species were collected biweekly from March through September 2011. Temporal variation of stable isotope compositions was observed in primary producers and consumer species. However, variation between replicates, particularly omnivores, often exceeded variation over time. The high degree of omnivory among the generalist feeders in this eelgrass community allows for generalist feeders to flexibly switch food sources, thus enhancing food-web stability. As coastal systems are subject to seasonal changes, as well as alterations related to human disturbance and climate, these food webs may retain a certain resilience due to their plentiful omnivores.

Beschreibung: Species identification based on morphological characteristics has caused misidentifications and led to twisted views of abundances and roles of ctenophores. Based on extensive field studies from 2007 to 2010, the occurrence of the arctic ctenophore Mertensia ovum was genetically verified in the southern, central and northern Baltic Sea, and its egg production, distribution and abundance were studied in relation to physical factors. Genetic analyses indicate that M. ovum is by far the most abundant small ctenophore in the Baltic Sea. Specimens from a 20 yr old ctenophore collection were also genetically identified as M. ovum, contrary to their previous morphological identification as another ctenophore species, Pleurobrachia pileus. Thus, earlier reports on P. pileus in the Baltic Sea may actually refer to M. ovum. The abundance of M. ovum was regulated by both salinity and temperature, with highest abundances found in sea areas and water layers at temperatures 〈7°C, salinities 〉5.5 and oxygen levels 〉4 ml l-1. During summer, the highest abundances of ctenophores and their eggs were found near the halocline, while the distribution was more uniform throughout the water column during winter. Only ctenophores 〉3.5 mm (oral-aboral length) produced eggs in the experiments, with an average rate of 2.2 eggs ind.-1 d-1. Finally, comparison with published data from the 1980s (assuming that those data refer to M. ovum) indicates that the present-day ctenophore abundance is ~80% lower in the north and ~55% higher in the southern parts of the Baltic Sea, due to reasons yet to be established.

Beschreibung: As part of the PeECE II mesocosm project, we investigated the effects of pCO2 levels on the initial step of heterotrophic carbon cycling in the surface ocean. The activities of microbial extracellular enzymes hydrolyzing 4 polysaccharides were measured during the development of a natural phytoplankton bloom under pCO2 conditions representing glacial (190 µatm) and future (750 µatm) atmospheric pCO2. We observed that (1) chondroitin hydrolysis was variable throughout the pre-, early- and late-bloom phases, (2) fucoidanase activity was measurable only in the glacial mesocosm as the bloom developed, (3) laminarinase activity was low and constant, and (4) xylanase activity declined as the bloom progressed. Concurrent measurements of microbial community composition, using denaturing-gradient gel electrophoresis (DGGE), showed that the 2 mesocosms diverged temporally, and from one another, especially in the late-bloom phase. Enzyme activities correlated with bloom phase and pCO2, suggesting functional as well as compositional changes in microbial communities in the different pCO2 environments. These changes, however, may be a response to temporal changes in the development of phytoplankton communities that differed with the pCO2 environment. We hypothesize that the phytoplankton communities produced dissolved organic carbon (DOC) differing in composition, a hypothesis supported by changing amino acid composition of the DOC, and that enzyme activities responded to changes in substrates. Enzyme activities observed under different pCO2 conditions likely reflect both genetic and population-level responses to changes occurring among multiple components of the microbial loop.

Beschreibung: Coral reefs are under threat, exerted by a number of interacting effects inherent to the present climate change, including ocean acidification and global warming. Bioerosion drives reef degradation by recycling carbonate skeletal material and is an important but understudied factor in this context. Twelve different combinations of pCO2 and temperature were applied to elucidate the consequences of ocean acidification and global warming on the physiological response and bioerosion rates of the zooxanthellate sponge Cliona orientalis—one of the most abundant and effective bioeroders on the Great Barrier Reef, Australia. Our results confirm a significant amplification of the sponges’ bioerosion capacity with increasing pCO2, which is expressed by more carbonate being chemically dissolved by etching. The health of the sponges and their photosymbionts was not affected by changes in pCO2, in contrast to temperature, which had significant negative impacts at higher levels. However, we could not conclusively explain the relationship between temperature and bioerosion rates, which were slightly reduced at both colder as well as warmer temperatures than ambient. The present findings on the effects of ocean acidification on chemical bioerosion, however, will have significant implications for predicting future reef carbonate budgets, as sponges often contribute the lion’s share of internal bioerosion on coral reefs.

Beschreibung: Epibiosis is one of the closest interspecies associations. The presence of epibionts potentially causes a multitude of beneficial or detrimental effects for the basibiont. It has been shown previously that large epibionts may increase the risk of dislodgement of bivalves. In this study, sublethal effects of epibiont-induced drag increase are investigated. I assessed (1) the effects of common epibiont species (Balanus improvisus, Enteromorpha intestinalis, Ectocarpus sp.) on drag properties of the host (the periwinkle Littorina littorea), and (2) the long-term consequences of drag increase on growth rates of snails living in steady flow. All epibiont species increase drag on the host snail. They do so to unequal extents. This may be due to morphological and hydrodynamic differences among the epibionts. Thus, per unit volume of epibiont, the filamentous alga Ectocarpus sp, has a substantially stronger effect than the barnacles. Synergistic effects on drag increase can be observed in a mixed aufwuchs community. As compared to clean conspecifics, snails bearing artificial epibionts grow 35% more slowly when exposed to moderate, steady flow (8 cm s(-1)) for 5 mo. This difference in growth rates is enhanced when food is limited. I hypothesize that fouled snails coping with higher drag invest more energy into foot activities (muscles and mucus). As a consequence, when food is limited, growth rates decrease in fouled snails.

Beschreibung: Pelagic zooplankton were monitored from 2000 to 2012 at a permanent location near the Svalbard archipelago, at the boundary between the central Arctic Ocean and the Greenland Sea in the eastern Fram Strait. The temporal results reveal the first evidence of successful reproduction in Arctic waters by an Atlantic pelagic crustacean from temperate waters. The Atlantic hyperid amphipod Themisto compressa is shown to have expanded its range from more southerly and warmer waters from 2004 onwards. Successful reproductive activity by T. compressa in Arctic waters was confirmed in 2011, indicated by the presence of a complete temporal series of developmental stages including ovigerous females and recently hatched juveniles. The Arctic amphipod community is currently in transition and a continuing northward spread of southern invaders could cause a biodiversity shift from large Arctic to smaller Atlantic species.

Beschreibung: To accurately assess community composition of invertebrates, both active and dormant life stages should be considered. Dormant stages are typically produced as a strategy to overcome inhospitable environmental conditions and can also facilitate species dispersal. While they often sink and accumulate in sediment of natural habitats forming ‘egg banks,’ dormant stages are also found in the sediments accumulated in ships’ ballast tanks. Recent studies have used 2 different methods to separate dormant stages from ballast sediment to assess invasion risk associated with ballast tanks: the colloidal silica sol Ludox HS 40 and sugar flotation (i.e. the Onbé-Marcus method). It has been assumed that the Ludox HS 40 method is most effective for separation but reduces dormant stage viability whereas sugar flotation has lower separation efficacy but higher resulting viability. We conducted a comparative assessment of the 2 methods by separating dormant stages from 160 ballast sediments and examining resulting abundance counts, hatching results, DNA extractions and PCR amplifications. We found no difference in the results between the methods. The financial cost of sugar flotation is lower than that of Ludox HS 40, and costs can be further reduced by using only 1 method instead of both due to lower labour costs, particularly for a large number of samples.

Beschreibung: Cyprinid herpesvirus 3 (CyHV-3) is the aetiological agent of a highly virulent and lethal disease of common carp Cyprinus carpio and its ornamental koi varieties. However, specific knowledge about immune mechanisms behind the infection process is very limited. We aimed to evaluate the effect of the CyHV-3 infection on the profile of 2 major components of the common carp immune acute phase response: the C-reactive protein (CRP) and the complement system. Common carp were infected with CyHV-3 by bath immersion. Fish were sampled before the infection and at 6, 12, 24, 72, 120 and 336 h post-infection for serum and head kidney, liver, gill and spleen tissues. CRP levels and complement activity were determined from the serum, whereas CRP- and complement-related genes (crp1, crp2, c1rs, bf/c2, c3, masp2) expression profiles were analysed in the tissues by quantitative PCR. Both CRP levels and complement activity increased significantly up to 10- and 3-fold, respectively, in the serum of infected fish during the challenge. Analysis revealed distinct organ- and time-dependent expression profile patterns for all selected genes. These results suggest that CRP and complement behave as acute phase reactants to CyHV-3 infection in common carp with an organ- and time-dependent response.

Beschreibung: The Kasatochi volcanic eruption that occurred in the central Aleutian Islands in Alaska, USA, in August 2008 is thought to have induced a massive diatom bloom in the iron-limited waters of the Gulf of Alaska, which potentially affected the oceanic food web by increasing the abundance of zooplankton and sockeye salmon Oncorhynchus nerka in the northeast Pacific Ocean. We report the first seawater experiments involving volcanic ash ejected from the Kasatochi eruption, showing that the ash released 61 to 83 nmol Fe, 374 to 410 nmol NO3-, 5 to 6 nmol PO43- and 170 to 585 nmol SiO2 when it contacted seawater. Our study suggests that the amount of iron released from Kasatochi ash (an increase of 2.0 to 2.8 nM Fe) was indeed sufficient to cause the observed phytoplankton bloom in the northeastern Pacific Gyre, while the impact of macronutrient release was minimal. We further evaluated the multiple, interdependent processes in the oceanic food web related to the diatom bloom, involving the ocean survival of juvenile salmon that entered the northeast Pacific Ocean in the summer of 2008.

Beschreibung: Concentrations of heme b, the ironcontaining prosthetic group of many hemoproteins, were measured in 6 species of marine phytoplankton (Dunaliella tertiolecta, Emiliania huxleyi, Thalassio - sira weissflogii, T. oceanica, Phaeodactylum tricor - nutum and Synechococcus sp. WH7803) that were subjected to variations in iron concentration. Changes in heme b in response to reduced light and nitrate were also ex amined for E. huxleyi and T. oceanica. Results from laboratory cultures were compared with heme b determined in particulate material in the North Atlantic. In cultures, heme b made up 18 ± 14% (SE) of the total iron pool. Reduced iron and nitrate concentrations resulted in a decreased intracellular heme b concentration, expressed as per mole carbon. Chlorophyll a (chl a) to heme b ratios in E. huxleyi and D. tertiolecta in creased in response to limited light and nutrient availability, but slightly decreased or did not change in the diatoms and the cyanophyte Synechococcus sp. WH7803. The heme b:particulate organic carbon (POC) and chl a:heme b ratios in the North Atlantic were within the range observed in phytoplankton cultures. In the surface mixed layer, decreases in heme b:POC ratios were linked to decreases in nutrient concentrations. Chl a:heme b ratios increased with depth and were thus primarily affected by light availability. Relative relationships between heme b, chl a and POC in the North Atlantic likely represented a change in the ability of cells to undertake cellular processes driven by chl a (light harvesting) and heme b (e.g. electron transport) according to ambient light and nutrient conditions.

Beschreibung: Stress often induces metabolically expensive countermeasures. Bivalve shell production is costly and can thus be indirectly impacted by environmental stress. Suboptimal salinity and temperature may constitute stressors that allocate energy away from shell production to cellular processes such as osmoregulation or to the repair of cellular damage. In the course of climate change, water temperatures of the Baltic Sea are predicted to increase, and salinity is predicted to regionally decrease. These shifts may lead to increased stress for temperate marine species adapted to relatively cool water temperatures and high salinity conditions. To better understand the importance of climate change-related stress, we assessed the isolated and interactive effects of salinity and temperature on shell increment (cumulative growth: shell), cellular oxidative stress (accumulation of oxidized lipids and proteins: lipofuscin), instantaneous physiological condition (condition index: CI), and mortality of young Mytilus edulis and Arctica islandica from the western Baltic Sea. Temperature and salinity interactively affected shell increment, lipofuscin accumulation, and mortality of M. edulis as well as shell increment of A. islandica. Shell increment of M. edulis was less affected by hyposalinity than shell increment of A. islandica. In both species the CI decreased and lipofuscin accumulation increased with increasing temperature. Lipofuscin accumulation negatively correlated with shell increment in M. edulis. We conclude that Baltic Sea populations of ecologically relevant bivalve species may experience severe stress by the predicted regional scenario of warming and desalination if evolutionary adaptation does not happen at a similar rate.

Beschreibung: Recent studies have discussed the consequences of ocean acidification for bacterial processes and diversity. However, the decomposition of complex substrates in marine environments, a key part of the flow of energy in ecosystems, is largely mediated by marine fungi. Although marine fungi have frequently been reported to prefer low pH levels, this group has been neglected in ocean acidification research. We present the first investigation of direct pH effects on marine fungal abundance and community structure. In microcosm experiments repeated in 2 consecutive years, we incubated natural North Sea water for 4 wk at in situ seawater pH (8.10 and 8.26), pH 7.82 and pH 7.67. Fungal abundance was determined by colony forming unit (cfu) counts, and fungal community structure was investigated by the culture-independent fingerprint method Fungal Automated Ribosomal Intergenic Spacer Analysis (F-ARISA). Furthermore, pH at the study site was determined over a yearly cycle. Fungal cfu were on average 9 times higher at pH 7.82 and 34 times higher at pH 7.67 compared to in situ seawater pH, and we observed fungal community shifts predominantly at pH 7.67. Currently, surface seawater pH at Helgoland Roads remains 〉8.0 throughout the year; thus we cannot exclude that fungal responses may differ in regions regularly experiencing lower pH values. However, our results suggest that under realistic levels of ocean acidification, marine fungi will reach greater importance in marine biogeochemical cycles. The rise of this group of organisms will affect a variety of biotic interactions in the sea.

Beschreibung: Since the Industrial Revolution, the partial pressure of carbon dioxide (pCO2) has been increasing and global ocean surface waters have absorbed 30% of the anthropogenic CO2 released into the atmosphere. An increase in pCO2 in surface ocean waters causes an increase in bicarbonate ions (HCO3-) and protons (H+) and a decrease in carbonate ions (CO32-), thereby decreasing the pH and the saturation state of the seawater with respect to CO32-. These changes in ocean chemistry (termed ocean acidification) are expected to have negative impacts on marine calcifying organisms. Because calcifying marine primary producers are important to the carbon cycle and rocky shore habitat structure and stability, investigating how they will respond to future oceanic pCO2 levels is a relevant and important topic of research. Due to a recent strong increase in the number of studies investigating the responses of calcifying marine macroalgae to elevated pCO2, this review aims to present the state of knowledge on the response of calcifying macroalgae to ocean acidification alone and in combination with global and local stressors. We discuss the physiological responses of calcifying macroalgae to elevated pCO2 within the contexts biogeography, taxonomy, and calcification mechanisms. Generally, coralline algae that deposit high-Mg calcite are most susceptible to high pCO2, and polar species are particularly at risk. However, some dolomite-depositing species may be able to acclimate to high pCO2. Calcifiers generally show sensitivity to overgrowth and outcompetition by noncalcifying algae when grown under elevated CO2 conditions, and this trend could be amplified under conditions of high inorganic nutrients. However, it still remains unknown whether or not calcifiers will be able to adapt to their rapidly changing environments. We discuss the lack of research on this topic, and provide some suggestions for how this knowledge gap can be filled by future research.

Beschreibung: Ocean acidification has the potential to affect growth and calcification of benthic marine invertebrates, particularly during their early life history. We exposed field-collected juveniles of Asterias rubens from Kiel Fjord (western Baltic Sea) to 3 seawater CO2 partial pressure (pCO2) levels (ranging from around 650 to 3500 µatm) in a long-term (39 wk) and a short-term (6 wk) experiment. In both experiments, survival and calcification were not affected by elevated pCO2. However, feeding rates decreased strongly with increasing pCO2, while aerobic metabolism and NH4+ excretion were not significantly affected by CO2 exposure. Consequently, high pCO2 reduced the scope for growth in A. rubens. Growth rates decreased substantially with increasing pCO2 and were reduced even at pCO2 levels occurring in the habitat today (e.g. during upwelling events). Sea stars were not able to acclimate to higher pCO2, and growth performance did not recover during the long-term experiment. Therefore, the top-down control exerted by this keystone species may be diminished during periods of high environmental pCO2 that already occur occasionally and will be even higher in the future. However, some individuals were able to grow at high rates even at high pCO2, indicating potential for rapid adaption. The selection of adapted specimens of A. rubens in this seasonally acidified habitat may lead to higher CO2 tolerance in adult sea stars of this population compared to the juvenile stage. Future studies need to address the synergistic effects of multiple stressors such as acidification, warming and reduced salinity, which will simultaneously impact the performance of sea stars in this habitat.

Beschreibung: The dynamics of methane production and oxidation in the sediments of Kiel Harbour (Baltic Sea) were studied over a period of 2 yr. Experimentally determined rates of methanogenesis varied between 13 and 82 mu mol CH4 m(-2) h(-1) integrated over the top 30 cm of sediment. The zone of maximum production was between 20 and 30 cm depth. High potential rates could be induced in sulfate-containing surface layers by the addition of molybdate, acetate or methanol, indicating competition with sulfate-reducing bacteria. The methane content in the sediment pore water increased from the surface to a seasonally varying depth of between 5 and 20 cm, where sulfate concentrations approached zero and methane concentrations of 3 to 4 mM were recorded. Aerobic methane oxidation rates in the surface sediment layer were in most cases higher than the methane flux expected to reach the sediment surface. Oxygen-depleted or anoxic conditions in the deep water, which frequently occurred during stratification periods in summer, resulted in methane gradients with increasing concentrations from the chemocline to the sediment. After recirculation of the water column in autumn in both years, unexpectedly high methane concentrations were also measured in the oxic water column of Kiel Harbour. This coincided with very reduced conditions in the surface sediment that allowed measurable methane production in the 0 to 10 cm layer.