ALBERT

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

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

Description: Chlorofluorocarbon-11 inventories for the deep Southern Ocean appear to confirm physical oceanographic and geochemical studies in the Southern Ocean, which suggest that no more than 5 × 106 cubic meters per second of ventilated deep water is currently being produced. This result conflicts with conclusions based on the distributions of the carbon-14/carbon ratio and a quasi-conservative property, PO4 *, in the deep sea, which seem to require an average of about 15 × 106cubic meters per second of Southern Ocean deep ventilation over about the past 800 years. A major reduction in Southern Ocean deep water production during the 20th century (from high rates during the Little Ice Age) may explain this apparent discordance. If this is true, a seesawing of deep water production between the northern Atlantic and Southern oceans may lie at the heart of the 1500-year ice-rafting cycle.

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

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

Description: A coral reef represents the net accumulation of calcium carbonate (CaCO3) produced by corals and other calcifying organisms. If calcification declines, then reef-building capacity also declines. Coral reef calcification depends on the saturation state of the carbonate mineral aragonite of surface waters. By the middle of the next century, an increased concentration of carbon dioxide will decrease the aragonite saturation state in the tropics by 30 percent and biogenic aragonite precipitation by 14 to 30 percent. Coral reefs are particularly threatened, because reef-building organisms secrete metastable forms of CaCO3, but the biogeochemical consequences on other calcifying marine ecosystems may be equally severe.

Description: Cephalopods play an important role in the trophic web of the Southern Ocean, but little information is available on their biology. The 2 largest sub-Antarctic seabirds, the king penguin Aptenodytes patagonicus and the wandering albatross Diomedea exulans, feed primarily on squids during the austral winter at the Crozet Islands. We examined a large number of accumulated cephalopod beaks in the stomach of these birds together with some undigested items; first, to understand how these 2 predators share the squid resource during winter, a period of supposed low food availability, and, second, to use a diving and a flying seabird as biological samplers of Southern Ocean cephalopods. Individuals of the family Onychoteuthidae formed the bulk of the squid diet, accounting for 72.6 and 57.0% of the number of lower beaks in samples from king penguins and wandering albatrosses, respectively. Seven different species were identified, the 3 main squids being Kondakovia longimana (38.8 and 28.0% by number for penguins and albatrosses, respectively), Moroteuthis ingens (13.5 and 26.2%) and M. knipovitchi (20.1 and 2.3%). Both seabirds preyed upon the same cephalopod species, but penguins primarily took small- to medium-sized juveniles (99.0% of the onychoteuthids) and albatrosses preyed on larger adult specimens (96.0%). Fresh remains indicated that adult K. longimana and M. ingens were mature individuals which, as shown by satellite tracking of albatrosses, were taken over the slope and nearby oceanic waters surrounding the archipelago. The present study indicates that mating/spawning of K. longimana and M. ingens occurs in Crozet waters during the winter months. It also extends the biogeography of K. longimana to north of the Antarctic Polar Front, in the Polar Frontal Zone, where it has not previously been recorded.

Description: The marine habitat exploited by black-browed Diomedea melanophrys and grey-headed albatrosses D. chrysostoma breeding at Campbell Island, New Zealand, was studied using satellite telemetry. Data were analysed in relation to the bathymetry and sea-surface temperature of the foraging zones. Black-browed albatrosses spent 55% of their time on the Campbell Plateau but also carried out long foraging trips to the Polar Front and Antarctic Zone at a distance of over 2000 km. They relied heavily on juvenile Micromesistius australis, a schooling fish, during foraging trips to the shelf but over oceanic waters the squid Martialia hyadesi was the main prey taken. Grey-headed albatrosses spent 71% of their time foraging over the deep waters of the Polar Frontal Zone where M. hyadesi comprised over 90% of the mass of prey taken. No satellite-tracked birds fed over the shelf, but data from the duration of foraging trips and dietary analysis suggests that shelf-feeding is important for this species. Significant inter-species differences in the time spent in neritic and oceanic zones show that black-browed albatrosses are reliant primarily on shelf resources while grey-headed albatrosses are primarily oceanic feeders. In addition, the 2 species overlapped little in the zones used over oceanic waters, with black-browed albatrosses feeding in more southerly waters than grey-headed albatrosses. However, both species feed on M. hyadesi when foraging in association with the Polar Front.

Description: The fishery for Illex argentinus in the Southwest Atlantic is subject to large inter-annual variability in recruitment strength. In this paper we attempt to build a predictive model using sea surface temperature (SST) to examine links between recruitment to the Falkland Islands fishery and environmental variability during the juvenile and adult life history stages. SST data from the National Center for Atmospheric Research (NCAR) were found to be comparable with near-surface data derived from in situ expendable bathy-thermograph (XBT) profiles in the southern Patagonian shelf. Variation in SST during the early life stages appears to be important in determining recruitment of I. argentinus. SST in the hatching grounds of the northern Patagonian shelf during the period of hatching (particularly June and July) was negatively correlated with catches in the fishery in the following season. SST anomaly data from positions in the Pacific and Southwest Atlantic were used to examine teleconnections between these areas. Links were seen at a lag of 2 yr between the Pacific and southern Patagonian shelf, and at about 5 yr between the Pacific and northern Patagonian shelf. This is consistent with SST anomalies associated with El Niño in the Pacific propagating around the globe via the Antarctic Circumpolar Wave (ACW). Predicting cold events via teleconnections between SST anomalies in the Pacific and Atlantic would appear to have the potential to predict the recruitment strength of I. argentinus in the Southwest Atlantic.

Description: Levels of genetic diversity and population differentiation were examined in temporally (1990 to 1997) and geographically separated samples of the argentine short-finned squid Illex argentinus using 7 microsatellite loci. Number of alleles (mean number of alleles per locus over all samples = 24.1) and heterozygosity (mean observed heterozygosity per sample = 0.84) were high for all samples, indicating that these loci have a greater potential utility for investigating population genetic structure than allozyme markers used in previous studies. Genetic diversity did not differ significantly between samples taken 5 yr after commencement of the fishery (1990) and those collected during a period of progressively intense fishing pressure (1994 and 1997). Several small but significant differences in between-sample genetic variation (FST) were observed, but these could not confirm the previous suggestion of cryptic species or several well-defined stocks within the fished population.

Description: Epibiosis is a spatially close association between 2 or more organisms belonging to the same or different species. Through direct and indirect interactions, this association has major effects on the species involved and on community dynamics. When the effects are predominantly beneficial for epibiont and basibiont, coevolution can be expected to lead to associational specificity. Circumstantial evidence, however, suggests that many epibionts are non-specific substratum-generalists. In this arti-cle, we investigate the commonness of specificity in epibiotic associations. In a first approach, we inves-tigated the in situ recruitment preferences of potential epibionts when choosing between artificial and living substrata. After exposure for 3 wk in early summer, an early successional community had estab-lished, comprising cyanobacteria, diatoms, sesslle colonial ciliates and red algae. All species recruited on almost all substrata available. However, artificial substrata were usually preferred over living sur-faces. Consequently, the species studied are class~fied as facultative epibionts. An analysis of a list of over 2000 epibiotic associations corroborated these results, the majority of described 'epibionts' are not basibiont-specific and generally occur on non-living substrata as well. Also, basibiont species usually bear more than 1 epibiont species. Relative to each other, epibionts and basibionts are characterised by a typical set of life history traits. We conclude that specific and obligate epibionts are rare. Their scarcity is discussed in view of multilevel antifouling defences and presumptive evolutionary transi-tions from epibiosis towards endoparasitism or endosymbiosis.

Description: Diving reptiles, unlike most diving birds and mammals, return infrequently to the surface to breathe. Spending the bulk of their lives underwater, they are likely to have developed a large variety of specific behavioural patterns different from those of their warm-blooded counterparts. However, for technical reasons, underwater behaviour of these aquatic reptiles remains poorly understood. In this study green turtles Chelonia mydas nesting on Cyprus (Eastern Mediterranean) were equipped with multi-channel data loggers monitoring diving behaviour and activity (via a logger-integrated 3-D compass which served as an activity sensor) during the inter-nesting interval. Data from 2 turtles for 2 consecutive inter-nesting intervals were available for detailed dive analysis. Both turtles showed highly variable dive patterns ranging from travelling subsurface dives to specific dive types such as U- (mainly resting and foraging dives), S- (a form of energy saving swimming) and V-dives. The green turtles stayed near the coast throughout the study, dived no deeper than ca 25 m, but remained underwater for up to ca 40 min. The recordings of the activity sensor revealed high activity levels (less than 20% resting d-1) during the whole inter-nesting period which was attributed to extensive foraging. The combination of both the activity data and the dive data showed that the turtles were engaged in travelling movements for 46% of the inter-nesting time spent underwater, foraged for 34% and rested for 12% of the time. We discuss the physiological, ecological and conservation implications of these results.

Description: Serial dilution experiments were conducted on JGOFS-North Atlantic cruise of RV 'Meteor' M36/2 at a 20° W transect in June and July 1996 to assess the role of microzooplankton grazing and nitrogen supply in controlling phytoplankton stocks in the subtropical and temperate northeast Atlantic. Rates of microzooplankton grazing ranged from 0.08 d-1 at 54° N to 0.53 d-1 at 40° N and mean growth rates of phytoplankton ranged from 0.19 d-1 at 54° N to 0.75 d-1 at 40° N. Both rates were positively related to seawater temperature, whereas the apparent growth yield of phytoplankton declined with increasing temperature from 0.19 µg chl a dm-3 d-1 at 54° N to 0.01 µg chl a dm-3 d-1 at 33° N. Complete nitrogen saturation of phytoplankton growth indicated light or non-nitrogenous limitation at the nitracline at 47° N and in the deep chlorophyll maximum at 33° N, whereas in the mixed layer at 47° N and 54° N the ambient nitrogen supply was sub-saturated and yielded 63 and 39% of nitrogen- saturated growth. Nitrogen supply of phytoplankton growth was dominated by external and cellular sources in nitrate-rich waters of the mixed layer at 54° N and at the nitracline at 47° N, whereas nitrogen regeneration dominated at the nitrate-depleted surface waters at 47° N. However, in the deep chlorophyll maxima at 33° N and 40° N phytoplankton growth was primarily maintained by nitrogen regeneration, although external nitrogen was sufficiently available. The recycling efficiency of the microbial community was defined as the ratio of regenerated growth yield to herbivorous grazing loss. Efficiencies of ~100% under post-bloom situations indicated tight coupling of predation, nitrogen supply and phytoplankton growth. We suggest that microzooplankton grazing has a high potential for nitrogen supply and biomass control of phytoplankton communities during summer in the temperate and subtropical northeast Atlantic.