ALBERT

Description: 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.

Description: 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.

Description: 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.

Description: 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.

Description: 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

Description: 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.

Description: 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.

Description: 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.

Description: 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.

Description: 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.

Description: The mechanisms involved in setting the annual cycle of the Florida Current transport are revisited using an adjoint model approach. Adjoint sensitivities of the Florida Current transport to wind stress reproduce a realistic seasonal cycle with an amplitude of ~1.2 Sv (1 Sv ≡ 106 m3 s−1). The annual cycle is predominantly determined by wind stress forcing and related coastal upwelling (downwelling) north of the Florida Strait along the shelf off the North American coast. Fast barotropic waves propagate these anomalies southward and reach the Florida Strait within a month, causing an amplitude of ~1 Sv. Long baroclinic planetary Rossby waves originating from the interior are responsible for an amplitude of ~0.8 Sv but have a different phase. The sensitivities corresponding to the first baroclinic mode propagate westward and are highly influenced by topography. Considerable sensitivities are only found west of the Mid-Atlantic Ridge, with maximum values at the western shelf edge. The second baroclinic mode also has an impact on the Florida Current variability, but only when a mean flow is present. A second-mode wave train propagates southwestward from the ocean bottom on the western side of the Mid-Atlantic Ridge between ~36° and 46°N and at Flemish Cap, where the mean flow interacts with topography, to the surface. Other processes such as baroclinic waves along the shelf and local forcing within the Florida Strait are of minor importance.

Description: The residual effect of surface gravity waves on mean flows in the upper ocean is investigated using thickness weighted mean (TWM) theory applied in a vertically Lagrangian and horizontally Eulerian coordinate system. Depth-dependent equations for the conservation of volume, momentum, and energy are derived. These equations allow for (i) finite amplitude fluid motions, (ii) the horizontal divergence of currents and (iii) a concise treatment of both the kinematic and viscous boundary conditions at the sea surface. Under the assumptions of steady and monochromatic waves and a uniform turbulent viscosity, the TWM momentum equations are used to illustrate the pressure- and viscosity-induced momentum fluxes through the surface that are implicit in previous studies of the wave-induced modification of the classical Ekman spiral problem. The TWM approach clarifies, in particular, the surface momentum flux associated with the so-called virtual wave stress of Longuet-Higgins. Overall the TWM framework can be regarded as an alternative to the three-dimensional Lagrangian mean framework of Pierson. Moreover the TWM framework can be used to include the residual effect of surface waves in large-scale circulation models. In specific models that carry the TWM velocity appropriate for advecting tracers as their velocity variable, the turbulent viscosity term should be modified so that the viscosity acts only on the Eulerian mean velocity.

Description: A method to derive salinity data from RAFOS float temperature and pressure measurements is described. It is based on evaluating the float's in situ density from its mechanical properties and in situ pressure and temperature data. The salinity of the surrounding water may then be determined, assuming that the float has reached equilibrium with its environment. This method, in comparison with the possible use of floatborne salinity cells, has the advantage of being both cost and energy neutral and highly stable in the long term. The effect on the estimated salinity of various parameters used in the determination of the float's in situ density is discussed. Results of seven RAFOS Boats deployed in the Brazil Basin are compared with corresponding CTD data to estimate the magnitude of these errors. At present, an accuracy of 0.3 psu is achieved. The accuracy may be improved to 0.02 psu by referring the float's calculated density to a reference density established by a CTD cast at the time of launch. Results from five floats deployed in the heterogeneous water masses of the Iberian Basin are compared with the corresponding CM casts to demonstrate the variability and interpretation of p-T-S float datasets from different areas.

Description: 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.

Description: Many aspects of barnacle body form are known to be developmentally plastic. Perhaps the most striking examples of such plasticity occur in their feeding legs and unusually long penises, the sizes and shapes of which can change dramatically and adaptively with changes in conspecific density and local water flow conditions. However, whether variation in overall appendage form is mirrored by structural responses in cuticle and muscle is not known. In order to determine how structural variation underlies phenotypic plasticity in barnacle appendages, we examined barnacles occurring at low and high population densities from one wave-protected and one wave-exposed site. We used histological sectioning and fluorescence microscopy of feeding legs and penises to compare cuticle thickness, muscle thickness, and muscle organization, and artificial penis inflation to compare penis extensibility. We observed striking differences in cuticle thickness, muscle thickness, and muscle organization between sites that differed in water velocity, but we found no clear differences associated with variation in conspecific density. Penis extensibility also did not differ consistently between sites. These results are consistent with an adaptive explanation for much of the remarkable and complex variation in barnacle feeding leg and penis morphology among sites that differ in water velocity.

Description: The Southern Hemisphere Subtropical Front (STF) is a narrow zone of transition between upper-level subtropical waters to the north and subantarctic waters to the south. It is found near 40 degrees S across the South Atlantic and South Indian Oceans and is associated with an eastward geostrophic current band, The current band in each basin is found at or just north of the surface front except near the eastern boundaries where most of the subtropical waters turn north into the eastern limbs of the subtropical gyres. The bands associated with the STF are thus distinct features separated from the strong zonal flows of the Antarctic Circumpolar Current farther south. The authors have referred to the current bands in the two respective oceans as the South Atlantic Current and the South Indian Ocean Current. In this paper the authors use the historical database from the South Pacific Ocean to investigate the geostrophic flow associated with the STF there. The STF extends across the southern Tasman Sea from south of Tasmania to southern New Zealand, and a weak eastward flow appears to be associated with it. The transport amounts to only about 3 Sv (1Sv = 10(6) m(3) s(-1)), little of which passes south of New Zealand. Mixing within the eddy-rich Tasman Sea may account for this weakness, while also setting up another more significant front in the northern Tasman Sea, the Tasman Front. It branches off from the East Australian Current toward the north of New Zealand, along which moves a flow of about 14 Sv. After passing north of New Zealand, a portion of this current flows east to contribute to a current band near 30 degrees S, while another portion turns south as the East Auckland Current and meets with subantarctic waters near Chatham Rise (44 degrees S), thus reestablishing the STF. An enhanced eastward current band is associated with the front there, one that extends across the remainder of the South Pacific and is referred to as the South Pacific Current. In comparison with its counterparts in the other basins, which typically begin by carrying 30 Sv (Atlantic) to 60 Sv (Indian) in the upper 1000 m in their western portions before weakening to 10-15 Sv in the east, the South Pacific Current is weak. Near Chatham Rise, it starts with a transport of approximately 5 Sv, and it remains near this strength as it shifts gradually north across the basin toward South America. The current appears to split into two smaller bands in the region of 115 degrees-85 degrees W, while near 28 degrees 5, 83 degrees W it begins to turn more strongly north and becomes shallower and weaker. Potential vorticity distributions indicate that this current acts as an impediment toward the northward spreading of Antarctic Intermediate Water, But why the South Pacific Current east of New Zealand should be so much weaker than its counterparts in the other basins is not particularly clear. It may be due to the presence of New Zealand and other topographic barriers to deep now east of Australia, to the axis of the subtropical gyre in the South Pacific shifting more rapidly southward with depth than those elsewhere, thus causing greater reductions in the underlying zonal velocities, and to strong poleward eddy heat and salt fluxes in the other two basins leading to smaller cross-STF gradients in the Pacific.

Description: 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.

Description: The suggestion is advanced that the remarkably low static stability of Antarctic surface waters may arise from a feedback loop involving global deep-water temperatures. If deep-water temperatures are too warm, this promotes Antarctic convection, thereby strengthening the inflow of Antarctic Bottom Water into the ocean interior and cooling the deep ocean. If deep waters are too cold, this promotes Antarctic stratification allowing the deep ocean to warm because of the input of North Atlantic Deep Water. A steady-state deep-water temperature is achieved such that the Antarctic surface can barely undergo convection. A two-box model is used to illustrate this feedback loop in its simplest expression and to develop basic concepts, such as the bounds on the operation of this loop. The model illustrates the possible dominating influence of Antarctic upwelling rate and Antarctic freshwater balance on global deep-water temperatures.

Description: 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

Description: 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.

Description: 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.

Description: The reliability of the Comprehensive Ocean–Atmosphere Dataset (COADS) Release 1a 2° monthly winds is tested by comparing it with instrumental measurements in the northwest Atlantic from 1981 to 1991. The instrumental dataset contains anemometer measurements of a very high homogeneity and quality, which were taken by six research sister ships with known anemometer heights in the northwest Atlantic. Special data processing was made with instrumental samples to provide compatibility with the COADS winds. Comparison shows overestimation of the COADS winds in the low ranges and underestimation of the strong and moderate winds. Application of the alternative equivalent Beaufort scales does not remove this bias and makes it even more pronounced. Thus, the conclusion is made that the disagreement obtained results primarily from the uncertainties of anemometer measurements in COADS, especially from the incorrect evaluation of the true wind. Instrumental data also do not indicate significant long-term interannual changes, which are pronounced in the COADS dataset for the 1980s. Some regional features of the comparison are discussed.

Description: 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.

Description: 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.

Description: 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.

Description: The eddy correlation technique is rapidly becoming an established method for resolving dissolved oxygen fluxes in natural aquatic systems. This direct and noninvasive determination of oxygen fluxes close to the sediment by simultaneously measuring the velocity and the dissolved oxygen fluctuations has considerable advantages compared to traditional methods. This paper describes the measurement principle and analyzes the spatial and temporal scales of those fluctuations as a function of turbulence levels. The magnitudes and spectral structure of the expected fluctuations provide the required sensor specifications and define practical boundary conditions for the eddy correlation instrumentation and its deployment. In addition, data analysis and spectral corrections are proposed for the usual nonideal conditions, such as the time shift between the sensor pair and the limited frequency response of the oxygen sensor. The consistency of the eddy correlation measurements in a riverine reservoir has been confirmed—observing a night–day transition from oxygen respiration to net oxygen production, ranging from −20 to +5 mmol m−2 day−1—by comparing two physically independent, eddy correlation instruments deployed side by side. The natural variability of the fluctuations calls for at least ∼1 h of flux data record to achieve a relative accuracy of better than ∼20%. Although various aspects still need improvement, eddy correlation is seen as a promising and soon-to-be widely applied method in natural waters.

Description: Some studies of ocean climate model experiments suggest that regional changes in dynamic sea level could provide a valuable indicator of trends in the strength of the Atlantic meridional overturning circulation (MOC). This paper describes the use of a sequence of global ocean–ice model experiments to show that the diagnosed patterns of sea surface height (SSH) anomalies associated with changes in the MOC in the North Atlantic (NA) depend critically on the time scales of interest. Model hindcast simulations for 1958–2004 reproduce the observed pattern of SSH variability with extrema occurring along the Gulf Stream (GS) and in the subpolar gyre (SPG), but they also show that the pattern is primarily related to the wind-driven variability of MOC and gyre circulation on interannual time scales; it is reflected also in the leading EOF of SSH variability over the NA Ocean, as described in previous studies. The pattern, however, is not useful as a “fingerprint” of longer-term changes in the MOC: as shown with a companion experiment, a multidecadal, gradual decline in the MOC [of 5 Sv (1 Sv ≡ 106 m3 s−1) over 5 decades] induces a much broader, basin-scale SSH rise over the mid-to-high-latitude NA, with amplitudes of 20 cm. The detectability of such a trend is low along the GS since low-frequency SSH changes are effectively masked here by strong variability on shorter time scales. More favorable signal-to-noise ratios are found in the SPG and the eastern NA, where a MOC trend of 0.1 Sv yr−1 would leave a significant imprint in SSH already after about 20 years.

Description: 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.

Description: Changes in the ventilation of the oxygen minimum zone (OMZ) of the tropical North Atlantic are studied using oceanographic data from 18 research cruises carried out between 28.5° and 23°W during 1999–2008 as well as historical data referring to the period 1972–85. In the core of the OMZ at about 400-m depth, a highly significant oxygen decrease of about 15 μmol kg−1 is found between the two periods. During the same time interval, the salinity at the oxygen minimum increased by about 0.1. Above the core of the OMZ, within the central water layer, oxygen decreased too, but salinity changed only slightly or even decreased. The scatter in the local oxygen–salinity relations decreased from the earlier to the later period suggesting a reduced filamentation due to mesoscale eddies and/or zonal jets acting on the background gradients. Here it is suggested that latitudinally alternating zonal jets with observed amplitudes of a few centimeters per second in the depth range of the OMZ contribute to the ventilation of the OMZ. A conceptual model of the ventilation of the OMZ is used to corroborate the hypothesis that changes in the strength of zonal jets affect mean oxygen levels in the OMZ. According to the model, a weakening of zonal jets, which is in general agreement with observed hydrographic evidences, is associated with a reduction of the mean oxygen levels that could significantly contribute to the observed deoxygenation of the North Atlantic OMZ.

Description: 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.

Description: The Rio Grande Rise acts as a natural barrier for the equatorward flow of Antarctic Bottom Water in the subtropical South Atlantic. In addition to the Vema Channel, the Hunter Channel cuts through this obstacle and offers a separate route for bottom-water import into the southern Brazil Basin. On the occasion of the Deep Basin Experiment, a component of the World Ocean Circulation Experiment (WOCE), the expected deep flow through the Hunter Channel was directly observed for the first time by an array of moored current meters and thermistor chains from December 1992 to May 1994. Main results are (i) the Hunter Channel is, in fact, a conduit for bottom-water flow into the Brazil Basin. Our new mean transport from moored current meters [2.92 (±1.24) × 106 m3 s−1] is significantly higher than earlier estimates that were based on geostrophic calculations. (ii) During the WOCE observational period a tendency toward increased bottom-water temperatures was observed. This observation from the Hunter Channel is consistent with findings from the Vema Channel. (iii) The overflow through the Hunter Channel is highly variable and puts in perspective earlier synoptic geostrophic transport estimates

Description: 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.

Description: The mean warm water transfer toward the equator along the western boundary of the South Atlantic is investigated, based on a number of ship surveys carried out during 1990–96 with CTD water mass observations and current profiling by shipboard and lowered (with the CTD/rosette) acoustic Doppler current profiler and with Pegasus current profiler. The bulk of the northward warm water flow follows the coast in the North Brazil Undercurrent (NBUC) from latitudes south of 10°S, carrying 23 Sv (Sv ≡ 106 m3 s−1) above 1000 m. Out of this, 16 Sv are waters warmer than 7°C that form the source waters of the Florida Current. Zonal inflow from the east by the South Equatorial Current enters the western boundary system dominantly north of 5°S, adding transport northwest of Cape San Roque, and transforming the NBUC along its way toward the equator into a surface-intensified current, the North Brazil Current (NBC). From the combination of moored arrays and shipboard sections just north of the equator along 44°W, the mean NBC transport was determined at 35 Sv with a small seasonal cycle amplitude of only about 3 Sv. The reason for the much larger near-equatorial northward warm water boundary current than what would be required to carry the northward heat transport are recirculations by the zonal current system and the existence of the shallow South Atlantic tropical–subtropical cell (STC). The STC connects the subduction zones of the eastern subtropics of both hemispheres via equatorward boundary undercurrents with the Equatorial Undercurrent (EUC), and the return flow is through upwelling and poleward Ekman transport. The persistent existence of a set of eastward thermocline and intermediate countercurrents on both sides of the equator was confirmed that recurred throughout the observations and carry ventilated waters from the boundary regime into the tropical interior. A strong westward current underneath the EUC, the Equatorial Intermediate Current, returns low-oxygen water westward. Consistent evidence for the existence of a seasonal variation in the warm water flow south of the equator could not be established, whereas significant seasonal variability of the boundary regime occurs north of the equator: northwestward alongshore throughflow of about 10 Sv of waters with properties from the Southern Hemisphere was found along the Guiana boundary in boreal spring when the North Equatorial Countercurrent is absent or even flowing westward, whereas during June–January the upper NBC is known to connect with the eastward North Equatorial Countercurrent through a retroflection zone that seasonally migrates up and down the coast and spawns eddies. The equatorial zone thus acts as a buffer and transformation zone for cross-equatorial exchanges, but knowledge of the detailed pathways in the interior including the involved diapycnal exchanges is still a problem.

Description: In this study, the authors discuss two different parameterizations for the effect of mixed layer eddies, one based on ageostrophic linear stability analysis (ALS) and the other one based on a scaling of the potential energy release by eddies (PER). Both parameterizations contradict each other in two aspects. First, they predict different functional relationships between the magnitude of the eddy fluxes and the Richardson number (Ri) related to the background state. Second, they also predict different vertical structure functions for the horizontal eddy fluxes. Numerical simulations for two different configurations and for a large range of different background conditions are used to evaluate the parameterizations. It turns out that PER is better suited to capture the Ri dependency of the magnitude of the eddy fluxes. On the other hand, the vertical structure of the meridional eddy fluxes predicted by ALS is more accurate than that of PER, while the vertical structure of the vertical eddy fluxes is well predicted by both parameterizations. Therefore, this study suggests the use of the magnitude of PER and the vertical structure functions of ALS for an improved parameterization of mixed layer eddy fluxes.

Description: 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.

Description: 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.

Description: 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.

Description: Northern Hemisphere stratospheric variability is investigated with respect to chaotic behavior using time series from three different variables extracted from four different reanalysis products and two numerical model runs with different forcing. The time series show red spectra at all frequencies and the probability distribution functions show persistent deviations from a Gaussian distribution. An exception is given by the numerical model forced with perpetual winter conditions—a case that shows more variability and follows a Gaussian distribution, suggesting that the deviation from Gaussianity found in the observations is due to the transition between summer and winter variability. To search for the presence of a chaotic attractor the correlation dimension and entropy, the Lyapunov spectrum, and the associated Kaplan–Yorke dimension are estimated. A finite value of the dimensions can be computed for each variable and data product, with the correlation dimension ranging between 3.0 and 4.0 and the Kaplan–Yorke dimension between 3.3 and 5.5. The correlation entropy varies between 0.6 and 1.1. The model runs show similar values for the correlation and Lyapunov dimensions for both the seasonally forced run and the perpetual-winter run, suggesting that the structure of a possible chaotic attractor is not determined by the seasonality in the forcing, but must be given by other mechanisms.

Description: 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.

Description: 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.

Description: Southern Hemisphere (SH) stratospheric variability is investigated with respect to chaotic behavior using time series from three different variables extracted from four different reanalysis products. The results are compared with the same analysis applied to the Northern Hemisphere (NH). The probability density functions (PDFs) for the SH show persistent deviations from a Gaussian distribution. The variability is given by white spectra for low frequencies, a slope of −1 for intermediate frequencies, and −3 slopes for high frequencies. Considering the time series for winter and summer separately, PDFs show a Gaussian distribution and the variability spectra change their slopes, indicating the role of the transition between winter and summer variability in shaping the time series. The correlation (D2) and the Kaplan–Yorke (DKY) dimensions are estimated. A finite value of the dimensions can be computed for each variable and data product, except for the NCEP zonal-mean zonal wind and temperature data, which violate the requirement D2 ≤ DKY, possibly owing to the presence of spurious trends and inconsistencies in the data. The value of D2 ranges between 2.6 and 3.9, while DKY ranges between 3.0 and 4.5. The results show that both D2 and DKY display large variability in their values both for different datasets and for different variables within the same dataset. The variability of the values of D2 and DKY thus leaves open the question about the existence of a low-dimensional attractor or if the finite dimensions of the system are the result of the projection of a larger attractor in a low-dimensional embedding space.

Description: In the present paper a hydrostatic “reduced gravity” model, generally used to simulate transient bottom-arrested gravity plumes, was coupled with a sediment transport model. The coupled model considers the respective contribution of suspended sediment particles on the buoyancy of a plume and allows one to simulate autosuspension and size-differential deposition of sediments based on the local turbulence and settling velocities. Simulations using the coupled model reveal that sediment-enriched plumes are able to inject both entrained and original shelf water masses into intermediate and bottom layers of an adjacent ocean basin in an ageostrophic dynamical balance. Hence the mechanism described here is more rapid than classic, “seawater” plumes, which are solely driven by surplus density of the water masses. Results suggest that “turbidity” plumes may constitute an important process in the formation and renewal of deep waters in the Arctic Ocean. In case a turbidity plume reaches its level of equilibrium density, deposition of suspended particles causes the density of the interstitial fluid to be lower than the density of the ambient fluid. This initiates upward convection within the water column. The substantial difference between TS- and turbidity plumes is described by model experiments that utilize idealized slope and sediment distributions. A realistic simulation of a turbidity plume cascading down the continental slope of the western Barents Sea is presented. The computed distribution of deposited sediments agrees well with observations in an area of high accumulation of shelf-derived sediments. The frequency of occurrence of sediment-enriched gravity plumes originating from the Barents Sea shelf is estimated from the various geological variables (thickness of sediments at the bottom, grain size composition) measured from bottom sediments samples.

Description: The Denmark Strait Overflow (DSO) supplies about one-third of the North Atlantic Deep Water and is critical to global thermohaline circulation. Knowledge of the pathways of DSO through the Irminger Basin and its transformation there is still incomplete, however. The authors deploy over 10 000 Lagrangian particles at the Denmark Strait in a high-resolution ocean model to study these issues. First, the particle trajectories show that the mean position and potential density of dense waters cascading over the Denmark Strait sill evolve consistently with hydrographic observations. These sill particles transit the Irminger Basin to the Spill Jet section (65.25°N) in 5–7 days and to the Angmagssalik section (63.5°N) in 2–3 weeks. Second, the dense water pathways on the continental shelf are consistent with observations and particles released on the shelf in the strait constitute a significant fraction of the dense water particles recorded at the Angmagssalik section within 60 days (~25%). Some particles circulate on the shelf for several weeks before they spill off the shelf break and join the overflow from the sill. Third, there are two places where the water density following particle trajectories decreases rapidly due to intense mixing: to the southwest of the sill and southwest of the Kangerdlugssuaq Trough on the continental slope. After transformation in these places, the overflow particles exhibit a wide range of densities.

Description: The sensitivities of the Brewer–Dobson circulation (BDC) to different distributions of tropical SST heating are investigated in an idealized aquaplanet model. It is found that an increase in tropical SSTs generally leads to an acceleration of tropical upwelling and an associated reduction in the age of air (AOA) in the polar stratosphere and that the AOA near the subtropical tropopause is correlated with local isentropic mixing of tropospheric air with stratospheric air. The zonal distribution of SST perturbations has a major impact on the vertical and meridional structure of the BDC as compared with other SST characteristics. Zonally localized SST heatings tend to generate a shallow acceleration of the stratospheric residual circulation, enhanced isentropic mixing associated with a weakened stratospheric jet, and a reduction in AOA mostly within the polar vortex. In contrast, SST heatings with a zonally symmetric structure tend to produce a deep strengthening of the stratospheric residual circulation, suppressed isentropic mixing associated with a stronger stratospheric jet, and a decrease of AOA in the entire stratosphere. The shallow versus deep strengthening of the stratospheric residual circulation change has been linked to wave propagation and dissipation in the subtropical lower stratosphere rather than wave generation in the troposphere, and the former can be strongly affected by the vertical position of the subtropical jet. These results suggest that, while the longitudinally localized SST trends under climate change may contribute to the change in the shallow branch of the BDC, the upward shift of the subtropical jet associated with the zonal SST heating can impact the deep branch of the BDC.

Description: The authors test the hypothesis that recent observed trends in surface westerlies in the Southern Hemisphere are directly consequent on observed trends in the timing of stratospheric final warming events. The analysis begins by verifying that final warming events have an impact on tropospheric circulation in a simplified GCM driven by specified equilibrium temperature distributions. Seasonal variations are imposed in the stratosphere only. The model produces qualitatively realistic final warming events whose influence extends down to the surface, much like what has been reported in observational analyses. The authors then go on to study observed trends in surface westerlies composited with respect to the date of final warming events. If the considered hypothesis were correct, these trends would appear to be much weaker when composited with respect to the date of the final warming events. The authors find that this is not the case, and accordingly they conclude that the observed surface changes cannot be attributed simply to this shift toward later final warming events.

Description: 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.

Description: 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.

Description: 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.

Description: The Atlantic meridional overturning circulation (AMOC) makes the strongest oceanic contribution to the meridional redistribution of heat. Here, an observation-based, forty-eight-month-long time series of the vertical structure and strength of the AMOC at 26.5°N is presented. From April 2004 to April 2008 the AMOC had a mean strength of 18.7 ±2.1 Sv with fluctuations of 4.8 Sv rms. The best guess of the peak-to-peak amplitude of the AMOC seasonal cycle is 6.7 Sv, with a maximum strength in autumn and a minimum in spring. While seasonality in the AMOC was commonly thought to be dominated by the northward Ekman transport, this study reveals that fluctuations of the geostrophic mid-ocean and Gulf Stream transports of 2.2 Sv and 1.7 Sv rms, respectively, are substantially larger than those of the Ekman component (1.2 Sv rms). A simple model based on linear dynamics suggests that the seasonal cycle is dominated by wind stress curl forcing at the eastern boundary of the Atlantic. Seasonal geostrophic AMOC anomalies might represent an important and previously underestimated component of meridional transport and storage of heat in the subtropical North Atlantic. There is evidence that the seasonal cycle observed here is representative of much longer intervals. Previously, hydrographic snapshot estimates between 1957 and 2004 had suggested a long-term decline of the AMOC by 8 Sv. This study suggests that aliasing of seasonal AMOC anomalies might have accounted for a large part of the inferred slowdown.

Description: 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.

Description: 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.

Description: 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.

Description: 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.

Description: Quantile mapping is routinely applied to correct biases of regional climate model simulations compared to observational data. If the observations are of similar resolution as the regional climate model, quantile mapping is a feasible approach. However, if the observations are of much higher resolution, quantile mapping also attempts to bridge this scale mismatch. Here, it is shown for daily precipitation that such quantile mapping-based downscaling is not feasible but introduces similar problems as inflation of perfect prognosis ("prog") downscaling: the spatial and temporal structure of the corrected time series is misrepresented, the drizzle effect for area means is overcorrected, area-mean extremes are overestimated, and trends are affected. To overcome these problems, stochastic bias correction is required.

Description: 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.

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

Description: A neural network (NN) has been developed in order to retrieve the cloud liquid water path (LWP) over the oceans from Special Sensor Microwave/Imager (SSM/I) data. The retrieval with NNs depends crucially on the SSM/I channels used as input and the number of hidden neurons—that is, the NN architecture. Three different combinations of the seven SSM/I channels have been tested. For all three methods an NN with five hidden neurons yields the best results. The NN-based LWP algorithms for SSM/I observations are intercompared with a standard regression algorithm. The calibration and validation of the retrieval algorithms are based on 2060 radiosonde observations over the global ocean. For each radiosonde profile the LWP is parameterized and the brightness temperatures (Tb’s) are simulated using a radiative transfer model. The best LWP algorithm (all SSM/I channels except T85V) shows a theoretical error of 0.009 kg m−2 for LWPs up to 2.8 kg m−2 and theoretical “clear-sky noise” (0.002 kg m−2), which has been reduced relative to the regression algorithm (0.031 kg m−2). Additionally, this new algorithm avoids the estimate of negative LWPs. An indirect validation and intercomparison is presented that is based upon SSM/I measurements (F-10) under clear-sky conditions, classified with independent IR-Meteosat data. The NN-based algorithms outperform the regression algorithm. The best LWP algorithm shows a clear-sky standard deviation of 0.006 kg m−2, a bias of 0.001 kg m−2, nonnegative LWPs, and no correlation with total precipitable water. The estimated accuracy for SSM/I observations and two of the proposed new LWP algorithms is 0.023 kg m−2 for LWP ⩽ 0.5 kg m−2.

Description: An initially resting ocean of stratification N is considered, subject to buoyancy loss at its surface of magnitude B0 over a circular region of radius r, at a latitude where the Coriolis parameter is f. Initially the buoyancy loss gives rise to upright convection as an ensemble of plumes penetrates the stratified ocean creating a vertically mixed layer. However, as deepening proceeds, horizontal density gradients at the edge of the forcing region support a geostrophic rim current, which develops growing meanders through baroclinic instability. Eventually finite-amplitude baroclinic eddies sweep stratified water into the convective region at the surface and transport convected water outward and away below, setting up a steady state in which lateral buoyancy flux offsets buoyancy loss at the surface. In this final state quasi-horizontal baroclinic eddy transfer dominates upright “plume” convection. By using “parcel theory” to consider the energy transformations taking place, it is shown that the depth, hfinal at which deepening by convective plumes is arrested by lateral buoyancy flux due to baroclinic eddies, and the time tfinal it takes to reach this depth, is given by both independent of rotation. Here γ and β are dimensionless constants that depend on the efficiency of baroclinic eddy transfer. A number of laboratory and numerical experiments are then inspected and carried out to seek confirmation of these parameter dependencies and obtain quantitative estimates of the constants. It is found that γ = 3.9 ± 0.9 and β = 12 ± 3. Finally, the implications of our study to the understanding of integral properties of deep and intermediate convection in the ocean are discussed.

Description: This paper describes, and establishes the dynamical mechanisms responsible for, the large-scale, time-mean, midlatitude circulation in a high-resolution model of the North Atlantic basin. The model solution is compared with recently proposed transport schemes and interpretations of the dynamical balances operating in the sub-tropical gyre. In particular, the question of the degree to which Sverdrup balance holds for the subtropical gyre is addressed. At 25°N, thermohaline-driven bottom flows cause strong local departures from the Sverdrup solution for the vertically integrated meridional mass transport, but these nearly integrate to zero across the interior of the basin. In the northwestern region of the subtropical gyre, in the vicinity of the Gulf Stream, higher-order dynamics become important, and linear vorticity dynamics is unable to explain the model's vertically integrated transport. In the subpolar gyre, the model transport bears little resemblance to the Sverdrup prediction, and higher-order dynamics are important across the entire longitudinal extent of the basin. The sensitivity of the model transport amplitudes, patterns, and dynamical balances are estimated by examining the solutions under a range of parameter choices and for four different wind stress forcing specifications. Taking into account a deficit of 7–10 Sv (Sv ≡ 106 m3 s−1) in the contribution of the model thermohaline circulation to the meridional transports at 25°N, the wind stress climatology of Isemer and Hasse appears to yield too strong of a circulation, while that derived from the NCAR Community Climate Model yields too weak of a circulation. The Hellerman and Rosenstein and ECMWF climatologies result in wind-driven transports close to observational estimates at 25°N. The range between cases for the annual mean southward transport in the interior above 1000 m is 14 Sv, which is 40%–70% of the mean transport itself. There is little sensitivity to the model closure parameters at this latitude. At 55°N, in the subpolar gyre, there is little sensitivity of the model solution to the choice of either closure parameters or wind climatology, despite large differences in the Sverdrup transports implied by the different wind stress datasets. Large year to year variability of the meridional transport east of the Bahamas makes it difficult to provide robust estimates of the sensitivity of the Antilles and deep western boundary current systems to forcing and parameter changes.

Description: A 12-month mooring record (May 1994–June 1995), together with accompanying PALACE float data, is used to describe an annual cycle of deep convection and restratification in the Labrador Sea. The mooring is located at 56.75°N, 52.5°W, near the former site of Ocean Weather Station Bravo, in water of 3500 m depth. This is a pilot experiment for climate monitoring, and also for studies of deep-convection dynamics. Mooring measurements include temperature (T), salinity (S), horizontal and vertical velocity, and acoustic measurement of surface winds. The floats made weekly temperature–salinity profiles between their drift level (near 1500 m) and the surface. With moderately strong cooling to the atmosphere (300 W m−2 averaged from November to March), wintertime convection penetrated from the surface to about 1750 m, overcoming the stabilizing effect of upper-ocean low-salinity water. The water column restratifies rapidly after brief vertical homogenization (in potential density, salinity, and potential temperature). Both the rapid restratification and the energetic high-frequency variations of T and S observed at the mooring are suggestive of a convection depth that varies greatly with location. Lateral variations in T and S exist down to very small scales, and these remnants of convection decay (with e-folding time 170 day) after convection ceases. Lateral variability at the scale of 100 km is verified by PALACE profiles. The Eulerian mooring effectively samples the convection in a mesoscale region of ocean as eddies sweep past it; the Lagrangian PALACE floats are complementary in sampling the geography of deep convection more widely. This laterally variable convection leaves the water column with significant vertical gradients most of the year. Convection followed by lateral mixing gives vertical salinity profiles the (misleading) appearance that a one-dimensional diffusive process is fluxing freshwater downward. During spring, summer, and fall the salinity, temperature, and buoyancy rise steadily with time throughout most of the water column. This is likely the result of mixing with the encircling boundary currents, compensating for the escape of Labrador Sea Water from the region. Low-salinity water mixes into the gyre only near the surface. The water-column heat balance is in satisfactory agreement with meteorological assimilation models. Directly observed subsurface calorimetry may be the more reliable indication of the annual-mean air–sea heat flux. Acoustic instrumentation on the mooring gave a surprisingly good time series of the vector surface wind. The three-dimensional velocity field consists of convective plumes of width 200 to 1000 m, vertical velocities of 2 to 8 cm s−1, and Rossby numbers of order unity, embedded in stronger (20 cm s−1) lateral currents associated with mesoscale eddies. Horizontal currents with timescales of several days to several months are strongly barotropic. They are suddenly energized as convection reaches great depth in early March, and develop toward a barotropic state, as also seen in models of convectively driven geostrophic turbulence in a weakly stratified, high-latitude ocean. Currents decay through the summer and autumn, apart from some persistent isolated eddies. These coherent, isolated, cold anticyclones carry cores of pure convected water long after the end of winter. Boundary currents nearby interact with the Labrador Sea gyre and provide an additional source of eddies in the interior Labrador Sea. An earlier study of the pulsation of the boundary currents is supported by observations of sudden ejection of floats from the central gyre into the boundary currents (and sudden ingestion of boundary current floats into the gyre interior), in what may be a mechanism for exchange between Labrador Sea Water and the World Ocean.

Description: A neural network is used to calculate the longwave net radiation (Lnet) at the sea surface from measurements of the Special Sensor Microwave/Imager (SSM/I). The neural network applied in this study is able to account largely for the nonlinearity between Lnet and the satellite-measured brightness temperatures (TB). The algorithm can be applied for instantaneous measurements over oceanic regions with the area extent of satellite passive microwave observations (30–60 km in diameter). Comparing with a linear regression method the neural network reduces the standard error for Lnet from 17 to 5 W m−2 when applied to model results. For clear-sky cases, a good agreement with an error of less than 5 W m−2 for Lnet between calculations from SSM/I observations and pyrgeometer measurements on the German research vessel Poseidon during the International Cirrus Experiment (ICE) 1989 is obtained. For cloudy cases, the comparison is problematic due to the inhomogenities of clouds and the low and different spatial resolutions of the SSM/I data. Global monthly mean values of Lnet for October 1989 are computed and compared to other sources. Differences are observed among the climatological values from previous studies by H.-J. Isemer and L. Hasse, the climatological values from R. Lindau and L. Hasse, the values of W. L. Darnell et al., and results from this study. Some structures of Lnet are similar for results from W. L. Darnell et al. and the present authors. The differences between both results are generally less than 15 W m−2. Over the North Atlantic Ocean the authors found a poleward increase for Lnet, which is contrary to the results of H.-J. Isemer and L. Hasse.

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

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

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

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

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

Description: The incidence angles of the SSM/I radiometers on the DMSP satellites vary from satellite to satellite and exhibit variations of up to 1.5° during one orbit. The effects of these variations on the measured brightness temperatures are investigated on the basis of simulated and measured data for oceanic arm. A deviation of 1° from the nominal incidence angle of 53.0° causes brightness temperature changes of up to 2 K depending on surface and atmospheric conditions. Errors of retrieved geophysical parameters on the order of 5%–10% result when the incidence angle variation is not taken into account. This is a common property of most published statistical algorithms. For total precipitable water and cloud liquid water content the error increases with increasing parameter value. For wind speed the error is largest for low wind speed and decreases with increasing wind speed. Due to the slowly varying latitudinal dependence of the incidence angle, these errors do not cancel out when monthly means are computed. A correction method is developed on the basis of simulated data and tested successfully with measured data. Observed brightness temperature differences between DMSP F10 and F11 are reduced when using corrected data. If diurnal variations of geophysical parameters are investigated, the incidence angle correction is mandatory to obtain useful results, especially for DMSP F10.

Description: In the autumn of 1996 the field component of an experiment designed to observe water mass transformation began in the Labrador Sea. Intense observations of ocean convection were taken in the following two winters. The purpose of the experiment was, by a combination of meteorological and oceanographic field observations, laboratory studies, theory, and modeling, to improve understanding of the convective process in the ocean and its representation in models. The dataset that has been gathered far exceeds previous efforts to observe the convective process anywhere in the ocean, both in its scope and range of techniques deployed. Combined with a comprehensive set of meteorological and air-sea flux measurements, it is giving unprecedented insights into the dynamics and thermodynamics of a closely coupled, semienclosed system known to have direct influence on the processes that control global climate.

Description: A dynamic–thermodynamic sea ice–mixed layer model for the Weddell Sea is complemented by a simple, diagnostic model to account for local sea ice–atmosphere interaction. To consider the atmospheric influence on the oceanic mixed layer, the pycnocline upwelling velocity is calculated using the theory of Ekman pumping. In several experiments, formation and conservation of a polynya in the Weddell Sea are investigated. Intrusion of heat into the lower atmosphere above the polynya area is assumed to cause a thermal perturbation and a cyclonic thermal wind field. Superposed with daily ECMWF surface winds, this modified atmospheric forcing field intensifies oceanic upwelling and induces divergent ice drift. Simulation results indicate that in case of a weak atmospheric cross-polynya flow the formation of a thermal wind field can significantly extend the lifetime of a large polynya. The repeated occurrence of the Weddell polynya in the years 1974–76 thus appears to be an effect of feedback mechanisms between sea ice, atmosphere, and oceanic mixed layer.

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

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

Description: Different processes have been proposed to explain the large-scale spreading of Mediterranean Water (MW) in the North Atlantic, however, no systematic study comparing the efficiency of different processes is yet available. Here, the authors present a series of experiments in a unified framework that is designed to quantify the effects of several physical processes on the spreading of MW in an idealized model of the North Atlantic. The common technique of restoring temperature and salinity to an observed distribution near the Mediterranean inflow fails to produce an adequate amount of MW because the eastern boundary region near the MW inflow is rather quiescent in models. Diapycnal processes like double diffusion and cabbeling turn out too inefficient to alone account for the large-scale MW anomaly. However, with a preexisting anomaly, double diffusion leads to a considerable northward and zonal redistribution of MW. The density anomaly induced by cabbeling curtails the zonal spreading of MW while it increases the northward spreading. With isopycnal mixing and the weak mean flow that prevails in the outflow region, a spatial distribution of the MW anomaly is obtained that is inconsistent with observations. Unrealistically high diffusion coefficients would be necessary to reproduce the observed salt flux into the Atlantic. The most effective process in the experiments is the volume flux associated with the Atlantic–Mediterranean exchange. The current system that is established in response to the inflow of MW into the Atlantic carries the anomaly almost 30° of longitude into the basin and along the eastern margin up to the northeastern corner of the domain and farther along the northern boundary.

Description: Parameterization of turbulent wind stress and sensible and latent heat fluxes is reviewed in the context of climate studies and model calculations, and specific formulas based on local measurements are recommended. Wind speed is of key importance, and in applying experimental results, the differences between local and modeled winds must be considered in terms of their method of observation or calculation. Climatological wind data based on Beaufort wind force reports require correction for historical trends. Integrated long-term net turbulent and radiative heat fluxes at the sea surface, calculated from archived data, are consistent with meridional heat transport through oceanographic sections; this lends support to the methods used

Description: The roles of ice particle size distributions (SDs) and particle shapes in cirrus cloud solar radiative transfer are investigated by analyzing SDs obtained from optical array probe measurements (particle sizes larger than 20–40 μm) during intensive field observations of the International Cirrus Experiment, the European Cloud and Radiation Experiment, the First ISCCP Regional Experiment, and the Central Equatorial Pacific Experiment. It is found that the cloud volume extinction coefficient is more strongly correlated with the total number density than with the effective particle size. Distribution-averaged mean single scattering properties are calculated for hexagonal columns, hexagonal plates, and polycrystals at a nonabsorbing (0.5 μm), moderately absorbing (1.6 μm), and strongly absorbing (3.0 μm) wavelength. At 0.5 μm (1.6 μm) (3.0 μm), the spread in the resulting mean asymmetry parameters due to different SDs is smaller than (comparable to) (smaller than) the difference caused by applying different particle shapes to these distributions. From a broadband solar radiative transfer point of view it appears more important to use the correct particle shapes than to average over the correct size distributions.

Description: A new optical disdrometer has been developed that is optimized for use in high wind speeds, for example, on board ships. The minimal detectable size of droplets is 0.35 mm. Each drop is measured separately with regard to its size and residence time within the sensitive volume. From the available information, the drop size distribution can be calculated with a resolution of 0.05 mm in diameter either by evaluation of the residence time of drops or by drop counting knowing the local wind. Experience shows that using the residence time leads to better results. Rain rates can be determined from the droplet spectra by assuming terminal fall velocity of the drops according to their size. Numerical modeling of disdrometer measurements has been performed, allowing the study of the effect of multiple occupancy of the sensitive volume and grazing incidences on disdrometer measurements. Based on these studies an iterative procedure has been developed to eliminate the impact of these effects on the calculated drop size distributions. This technique may also be applied to any other kind of disdrometer. Long-term simultaneous measurements of the disdrometer and a conventional rain gauge have been used to validate this procedure.

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

Description: The compatibility of the Gent and McWilliams thickness mixing parameterization with perturbation thickness fluxes evaluated from eddy-resolving North Atlantic model results is investigated. After extensive spatial and temporal averaging, a linear correlation between the parameterized fluxes and those calculated directly from model fluctuations in the subtropics could be found. A direct estimate of a constant mixing parameter κ could be inferred in the order of 1.0 × 107 cm2 s−1.

Description: A simple point-vortex “heton” model is used to study localized ocean convection. In particular, the statistically steady state that is established when lateral buoyancy transfer, effected by baroclinic instability, offsets the localized surface buoyancy loss is investigated. Properties of the steady state, such as the statistically steady density anomaly of the convection region, are predicted using the hypothesis of a balance between baroclinic eddy transfer and the localized surface buoyancy loss. These predictions compare favorably with the values obtained through numerical integration of the heton model. The steady state of the heron model can be related to that in other convection scenarios considered in several recent studies by means of a generalized description of the localized convection. This leads to predictions of the equilibrium density anomalies in these scenarios, which concur with those obtained by other authors. Advantages of the heton model include its inviscid nature, emphasizing the independence of the fluxes affected by the baroclinic eddies from molecular processes, and its extreme economy, allowing a very large parameter space to be covered. This economy allows us to examine more complicated forcing scenarios: for example, forcing regions of varying shape. By increasing the ellipticity of the forcing region, the instability is modified by the shape and, as a result, no increase in lateral fluxes occurs despite the increased perimeter length. The parameterization of convective mixing by a redistribution of potential vorticity, implicit in the heton model, is corroborated; the heton model equilibrium state has analogous quantitative scaling behavior to that in models or laboratory experiments that resolve the vertical motions. The simplified dynamics of the heton model therefore allows the adiabatic advection resulting from baroclinic instability to be examined in isolation from vertical mixing and diffusive processes. These results demonstrate the importance of baroclinic instability in controlling the properties of a water mass generated by localized ocean convection. A complete parameterization of this process must therefore account for the fluxes induced by horizontal variations in surface buoyancy loss and affected by baroclinic instability.

Description: On the basis of the collection of individual marine observations available from the Comprehensive Ocean–Atmosphere Data Set, major parameters of the sea state were evaluated. Climatological fields of wind waves and swell height and period, as well as significant wave height and resultant period are obtained for the North Atlantic Ocean for the period from 1964 to 1993. Validation of the results against instrumental records from National Data Buoy Center buoys and ocean weather station measurements indicate relatively good agreement for wave height and systematic biases in the visually estimated periods that were corrected. Wave age, which is important for wind stress estimates, was evaluated form wave and wind observations. The climatology of wave age indicates younger waves in winter in the North Atlantic midlatitudes and Tropics. Wave age estimates were applied to the calculations of the wind stress using parameterizations from field experiments. Differences between wave-age-based and traditional estimates are not negligible in wintertime in midlatitudes and Tropics where wave-induced stress contributes from 5% to 15% to the total stress estimates. Importance of the obtained effects for ocean circulation and climate variability is discussed.

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

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

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

Description: Analysis of modern and historical observations demonstrates that the temperature of the intermediate-depth (150–900 m) Atlantic water (AW) of the Arctic Ocean has increased in recent decades. The AW warming has been uneven in time; a local 1°C maximum was observed in the mid-1990s, followed by an intervening minimum and an additional warming that culminated in 2007 with temperatures higher than in the 1990s by 0.24°C. Relative to climatology from all data prior to 1999, the most extreme 2007 temperature anomalies of up to 1°C and higher were observed in the Eurasian and Makarov Basins. The AW warming was associated with a substantial (up to 75–90 m) shoaling of the upper AW boundary in the central Arctic Ocean and weakening of the Eurasian Basin upper-ocean stratification. Taken together, these observations suggest that the changes in the Eurasian Basin facilitated greater upward transfer of AW heat to the ocean surface layer. Available limited observations and results from a 1D ocean column model support this surmised upward spread of AW heat through the Eurasian Basin halocline. Experiments with a 3D coupled ice–ocean model in turn suggest a loss of 28–35 cm of ice thickness after 50 yr in response to the 0.5 W m−2 increase in AW ocean heat flux suggested by the 1D model. This amount of thinning is comparable to the 29 cm of ice thickness loss due to local atmospheric thermodynamic forcing estimated from observations of fast-ice thickness decline. The implication is that AW warming helped precondition the polar ice cap for the extreme ice loss observed in recent years.

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

Description: The importance of decadal climate variability (DCV) research is being increasingly recognized, including by the World Climate Research Program (WCRP) and the Intergovernmental Panel on Climate Change (IPCC). An improved understanding of DCV is very important because stakeholders and policymakers want to know the likely climate trajectory for the coming decades for applications to water resources, agriculture, energy, and infrastructure development. Responding to this demand, many climate modeling groups in the United States, Europe, Japan, and elsewhere are gearing up to assess the potential for decadal climate predictions. The magnitudes of regional DCV often exceed those associated with the trends resulting from anthropogenic changes. Therefore, differentiating between the two is also very important for planning, implementation, and national and international treaties.

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

Description: In the Gulf of Lions, observations of deep convection have been sporadically carried out over the past three decades, showing significant interannual variability of convection activity. As long time series of meteorological observations of the region are available from coastal stations, heat flux time series for the Gulf of Lions for the individual winters from 1969 to 1994 are derived by calibrating these observations against direct measurements obtained over the convection site. These heat fluxes are also compared against heat fluxes obtained by the French PERIDOT weather model for the winter of 1991/92. A Kraus–Turner one-dimensional mixed layer model is initialized by climatological mean temperature and salinity profiles and then driven by the heat flux time series of the individual years. Resulting convection depths are in satisfactory agreement with existing observational evidence, showing the dominance of interannual variability of local forcing on convection variability. The interannual variability of convection depth causes interannual variations in deep-water properties, and these are also compared with the hydrographic database.

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

Description: Previous attempts to derive the depth-dependent expression of the radiation stress have lead to a debate concerning (i) the applicability of Mellor’s approach to a sloping bottom, (ii) the introduction of the delta function at the mean sea surface in the later papers by Mellor, and (iii) a wave-induced pressure term derived in several recent studies. The authors use an equation system in vertically Lagrangian and horizontally Eulerian (VL) coordinates suitable for a concise treatment of the surface boundary, and obtain an expression for the depth-dependent radiation stress that is consistent with the vertically-integrated expression given by Longuet-Higgins and Stewart. Concerning (i)-(iii) in the above, the difficulty of handling a sloping bottom disappears when wave-averaged momentum equations in the VL coordinates are written for the development of (not the Lagrangian mean velocity but) the Eulerian mean velocity. There is also no delta function at the sea surface in the expression for the depth-dependent radiation stress. The connection between the wave-induced pressure term in the recent studies and the depth-dependent radiation stress term is easily shown by rewriting the pressure-based form stress term in the thickness-weighted-mean (TWM) momentum equations as a velocity-based term which contains the time derivative of the pseudomomentum in the TWM framework.

Description: The upper ocean, including the biologically productive euphotic zone and the mixed layer, has great relevance for studies of physical, biogeochemical, and ecosystem processes and their interaction. Observing this layer with a continuous presence, sampling many of the relevant variables, and with sufficient vertical resolution, has remained a challenge. Here a system is presented which can be deployed on the top of deep-ocean moorings, with a drive mechanism at depths of 150-200m, which mechanically winches a large sensor float and smaller communications float tethered above it to the surface and back down again, typically twice per day for periods up to 1 year. The sensor float can carry several sizeable sensors, and it has enough buoyancy to reach the near surface and for the communications float to pierce the surface even in the presence of strong currents. The system can survive mooring blow-over to 1000m depth. The battery-powered design is made possible by using a balanced energy-conserving principle. Reliability is enhanced with a drive assembly that employs a single rotating part that has no slip rings or rotating seals. The profiling bodies can break the surface to sample the near-surface layer and to establish satellite communication for data relay or reception of new commands. An inductive pass-through mode allows communication with other mooring components throughout the water column beneath the system. A number of successful demonstration deployments have been completed.

Description: In recent years, profiling floats, which form the basis of the successful international Argo observatory, are also being considered as platforms for marine biogeochemical research. This study showcases the utility of floats as a novel tool for combined gas measurements of CO2 partial pressure (pCO2) and O2. These float prototypes were equipped with a small-sized and submersible pCO2 sensor and an optode O2 sensor for high resolution measurements in the surface ocean layer. Four consecutive deployments were carried out during Nov. 2010 and June 2011 near the Cape Verde Ocean Observatory (CVOO) in the eastern tropical North Atlantic. The profiling float performed upcasts every 31 h while measuring pCO2, O2, salinity, temperature and hydrostatic pressure in the upper 200 m of the water column. In order to maintain accuracy, regular pCO2 sensor zeroings at depth and surface, as well as optode measurements in air, were performed for each profile. Through the application of data processing procedures (e.g., time-lag correction) accuracies of float-borne pCO2 measurements were greatly improved (10 – 15 μatm for water column and 5 μatm for surface measurements). O2 measurements yielded an accuracy of 2 μmol kg−1. First results of this pilot study show the possibility of using profiling floats as a platform for detailed and unattended observations of the marine carbon and oxygen cycle dynamics.

Description: Simple idealized layered models and primitive equation models show that the meridional gradient of the zonally averaged pressure has no direct relation with the meridional flow. This demonstrates a contradiction in an often-used parameterization in zonally averaged models. The failure of this parameterization reflects the inconsistency between the model of Stommel and Arons and the box model of Stommel, as previously pointed out by Straub. A new closure is proposed. The ocean is divided in two dynamically different regimes: a narrow western boundary layer and an interior ocean; zonally averaged quantities over these regions are considered. In the averaged equations three unknowns appear: the interior zonal pressure difference Delta p(i), the zonal pressure difference Delta p(b) of the boundary layer, and the zonal velocity us at the interface between the two regions. Here Delta p(i) is parameterized using a frictionless vorticity balance, Delta p(b), by the difference of the mean pressure in the interior and western boundary, and u(delta) by the mean zonal velocity of the western boundary layer. Zonally resolved models, a layer model, and a primitive equation model validate the new parameterization by comparing with the respective zonally averaged counterparts. It turns out that the zonally averaged models reproduce well the buoyancy distribution and the meridional flow in the zonally resolved model versions with respect to the mean and time changes.

Description: The use of a coupled ocean/atmosphere/sea-ice model to hindcast (i.e. historical forecast) recent climate variability is described and illustrated for the cases of the 1976/77 and 1998/99 climate shift events in the Pacific. The initialization is achieved by running the coupled model in partially coupled mode whereby global observed wind stress anomalies are used to drive the ocean/sea-ice component of the coupled model while maintaining the thermodynamic coupling between the ocean/sea-ice and atmosphere components. Here we show that hindcast experiments can successfully capture many features associated with the 1976/77 and 1998/99 climate shifts. For instance, hindcast experiments started from the beginning of 1976 can capture sea surface temperature (SST) warming in the central-eastern equatorial Pacific and the positive phase of the Pacific Decadal Oscillation (PDO) throughout the 9 years following the 1976/77 climate shift, including the deepening of the Aleutian low pressure system. Hindcast experiments started from the beginning of 1998 can also capture part of the anomalous conditions during the 4 years after the 1998/99 climate. We argue that the dynamical adjustment of heat content anomalies that are present in the initial conditions in the tropics is important for the successful hindcast of the two climate shifts.

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

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

Description: The Cape Verde Frontal Zone separates the North and the South Atlantic Central Waters in the eastern North Atlantic. It also represents the boundary between the ventilated subtropical gyre and the quasi-stagnant shadow zone in the southeast. The thermohaline front is nearly compensated with respect to density, and density parameters RP, suggest the existence of double-diffusive processes. Datasets from three cruises to the region, approximately one year apart each, are used to determine the effects of double-diffusive diapycnal versus isopycnal mixing. For this purpose results from the usual temperature-salinity analysis assuming isopycnal mixing are compared to results from a multiparameter analysis where nutrient and oxygen data are also used. Significant diapycnal fluxes are found in the frontal zone between 200 and 300 m, with water mass contents being changed by more than 20% through diapycnal mixing. The associated buoyancy fluxes have a similar magnitude as surface fluxes in the area and thus represent an important contribution to the vertical balances of heat and salt.

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