Escape responses of copepod nauplii in the flow field of the blue mussel,<Emphasis Type="Italic"> Mytilus edulis</Emphasis>

Bivalves are important grazers on phytoplankton in shallow waters. However, very little is known about their ability to capture actively moving zooplankton. We investigated the escape response and success of early and late nauplii of three copepod species (Acartia tonsa, Temora longicornis and Eurytemora affinis) in the flow field of a blue mussel, Mytilus edulis, using both video observations and incubation experiments. An empirical model was created to describe the spatial distribution of the fluid deformation rate. Nauplii responded with escape jumps at mean fluid deformation rates of 0.6-1.9 s^у. Escape success differed between taxa. T. longicornis was the poorest escaper, while A. tonsa and E. affinis were more efficient and similar to one another. Deformation rates differed in different parts of the flow field, which resulted in differences in escape success between the sectors. Nauplii were caught most often in the sector furthest away from the exhalent siphon, where the deformation rate was the weakest. There the nauplii were unable to detect an escape signal in time to react and flee.     

Characterization of urease activity in three marine phytoplankton species,<Emphasis Type="Italic"> Aureococcus anophagefferens</Emphasis>,<Emphasis Type="Italic"> Prorocentrum minimum</Emphasis>, and<Emphasis Type="Italic"> Thalassiosira weissflogii</Emphasis>

The availability of different forms of nitrogen in coastal and estuarine waters may be important in determining the abundance and productivity of different phytoplankton species. Although urea has been shown to contribute as much as 50% of the nitrogen for phytoplankton nutrition, relatively little is known of the activity and expression of urease in phytoplankton. Using an in vitro enzyme assay, urease activities were examined in laboratory cultures of three species: Aureococcus anophagefferens Hargraves et Sieburth, Prorocentrum minimum (Pavillard) Schiller, and Thalassiosira weissflogii (Grunow) Fryxell et Hasle. Cultures of P. minimum and T. weissflogii were grown on three nitrogen sources (NO₃^m, NH₄⁺, and urea), while A. anophagefferens was grown only on NO₃^m and urea. Urease was found to be constitutive in all cultures, but activity varied with growth rate and assay temperature for the different cultures. For A. anophagefferens, urease activity varied positively with growth rate regardless of the N source, while for P. minimum, urease activity varied positively with growth rate only for cultures grown on urea and NH₄⁺. In contrast, for T. weissflogii, activity did not vary with growth rate for any of the N sources. For all species, urease activity increased with assay temperature, but with different apparent temperature optima. For A. anophagefferens, in vitro activity increased from near 0-30°C, and remained stable to 50°C, while for P. minimum, increased in vitro activity was noted from near 0-20°C, but constant activity was observed between 20°C and 50°C. For T. weissfloggii, while activity also increased from 0°C to 20°C, subsequent decreases were noted when temperature was elevated above 20°C. Urease activity had a half-saturation constant of 120-165 wg atom N l^у in all three species. On both an hourly and daily basis, urease activity in A. anophagefferens exceeded nitrogen demand for growth. In P. minimum, urease activity on an hourly basis matched the nitrogen demand, but was less than the demand on a daily basis. For T. weissflogii, urease activity was always less than the nitrogen demand. These patterns in urease activity in three different species demonstrate that while apparently constitutive, the regulation of activity was substantially different in the diatom. These differences in the physiological regulation of urease activity, as well as other enzymes, may play a role in their ecological success in different environments.     

Evaluation of fatty acids as biomarkers for a natural plankton community. A field study of a spring bloom and a post-bloom period off West Greenland

An investigation of the fatty acid composition of a natural arctic plankton community was carried out over two fishing banks located between 63°N and 65°N off the West Greenland coast. Samples for fatty acid analyses, species determination and biomass assessments of the plankton community were taken at the depth of fluorescence maximum. High biomass and diatom dominance during the spring bloom and low biomass and flagellate dominance in the post-bloom period were reflected by the fatty acid profiles. The total amount of fatty acid ranged from 55 to 132 µg l^-1 during the spring bloom and from 1 to 5 µg l^-1 during the post bloom. Analysis of the fatty acids showed that when the plankton was dominated by diatoms of the genera Thalassiosira and Chaetoceros, the proportions of C16:1(n-7) and C20:5(n-3) were correspondingly high. C18s, and particularly C18:1(n-9), were more abundant when the plankton was dominated by small autotrophic flagellates, primarily haptophytes. We found a good positive correlation between the common diatom marker, C16:1(n-7)/C16:0, and the biomass percentage of diatoms (r=0.742, P<0.001), as well as between the biomass percentage of flagellates and total C18 fatty acids (r=0.739, P<0.001). This supports the use of these specific fatty acids and fatty acid ratios as general biomarkers of the plankton community. However, the fatty acids are not specific enough to sufficiently characterise the composition of the plankton community, and microscopical support is needed to verify observed trends.     

Large-scale oceanic distribution and population structure of Calanus finmarchicus, in relation to physical environment, food and predators

The investigation was carried out from 62°N to 73°N and from 14°E to 11°W in the Norwegian Sea during 19 June-12 July 1997. Regional differences in the phase of the seasonal development of the plankton community were evident, most pronounced across the Arctic front. In the Coastal and eastern Atlantic domains, post-bloom conditions prevailed, characterised by low chlorophyll a (chl a) levels and a phytoplankton assemblage dominated by coccolithophorids and small flagellates. During the study period, egg production rates of Calanus finmarchicus were low (<10 eggs female^-1 day^-1), older copepodite stages dominated, and the seasonal descent to deeper waters had started. In the Arctic domain, bloom conditions were evident by high chl a levels and a high abundance of large diatoms. Egg production rates were higher (a maximum of 29 eggs female^-1 day^-1), but the dominance of stages CI-CIII indicated that considerable spawning had already occurred prior to the spring bloom. The seasonal descent had barely started. Both invertebrate and fish predators were most abundant in the Coastal and eastern Atlantic domains, with abundance strongly decreasing north-westwards. No tight relationship between total abundance of invertebrate or fish predators and that of C. finmarchicus was apparent. However, a weak, but significant, relationship between abundance of young stages of chaetognaths and Euchaeta spp. versus young stages of C. finmarchicus was found, indicating that these invertebrate predators develop parallel to the development of the new cohort of C. finmarchicus. In early summer, C. finmarchicus had reached overwintering stages, and had started to accumulate in deeper waters in areas with the highest abundance of horizontally migratory planktivorous fish.     

Cross-shelf variation in calanoid copepod production during summer 1996 off the Oregon coast, USA

The fecundity of nine species of adult female calanoid copepods, and molting rates for copepodite stages of Calanus marshallae were measured in 24 h shipboard incubations from samples taken during the upwelling season off the Oregon coast. Hydrographic and chlorophyll measurements were made at approximately 300 stations, and living zooplankton were collected at 36 stations on the continental shelf (<150 m depth) and 37 stations offshore of the shelf (>150 m depth) for experimental work. In our experiments, maximum egg production rates (EPR) were observed only for Calanus pacificus and Pseudocalanus mimus, 65.7 and 3.9 eggs fem^-1 day^-1 respectively, about 95% of the maximum rates known from published laboratory observations. EPR of all other copepod species (e.g., C. marshallae, Acartia longiremis and Eucalanus californicus) ranged from 3% to 65% of maximum published rates. Fecundity was not significantly related to body weight or temperature, but was significantly correlated with chlorophyll a concentration for all species except Paracalanus parvus and A. longiremis. Copepod biomass and production in on-shelf waters was dominated by female P. mimus and C. marshallae, accounting for 93% of the adult biomass (3.1 mg C m^-3) and 81% of the adult production (0.19 mg C m^-3 day^-1). Biomass in the off-shelf environment was dominated by female E. californicus, P. mimus, and C. pacificus, accounting for 95% of the adult biomass (2.2 mg C m^-3) and 95% of the adult production (0.08 mg C m^-3 day^-1). Copepodite (C1-C5) production was estimated to be 2.1 mg C m^-3 day^-1 (on-shelf waters) and 1.2 mg C m^-3 day^-1 (off-shelf water). Total adult + juvenile production averaged 2.3 mg C m^-3 day^-1 (on-shelf waters) and 1.3 mg C m^-3 day^-1 (off-shelf waters). We compared our measured female weight-specific growth rates to those predicted from the empirical models of copepod growth rates of Huntley and Lopez [Am Nat (1992) 140:201-242] and Hirst and Lampitt [Mar Biol (1998) 132:247-257]. Most of our measured values were lower than those predicted from the equation of Huntley and Lopez. We found good agreement with Hirst and Lampitt for growth rates <0.10 day^-1 but found that their empirical equations underestimated growth at rates >0.10 day^-1. The mismatch with Hirst and Lampitt resulted because some of our species were growing at maximum rates whereas their composite empirical equations predict "global" averages that do not represent maximum growth rates.     

In situ feeding rates and grazing impact of zooplankton in a South African temporarily open estuary

Recent studies in temporarily open estuaries of South Africa have shown that phytoplankton biomass is at times low, when compared to the high standing stock of the grazers. In situ grazing rates of the dominant zooplankton species were estimated at the Mpenjati Estuary once during the winter closed phase, in August 1999, and once during the summer open phase, in February 2000. The study aimed at determining what proportion of the energetic demands of the dominant grazers of the estuary is met by the available phytoplankton. Results show that the gut of all species exhibited higher pigment concentrations during the night than during the day, both in winter and summer. Gut pigment contents ranged from 0.27 to 5.38 ng pigm individual^-1 in the mysid Gastrosaccus brevifissura, from 0.16 to 1.63 ng pigm individual^-1 in the copepod Pseudodiaptomus hessei, from 0.12 to 0.45 ng pigm individual^-1 in the copepod Acartia natalensis, and from 0.8 to 5.44 ng pigm individual^-1 in the caridean Palaemon sp. [where pigm is the sum of chlorophyll-a (chl-a) and phaeopigments]. During the winter closed phase, gut evacuation rates for G. brevifissura, P. hessei, and A. natalensis were 0.62, 0.42, and 0.46 h^-1, respectively. In summer, gut evacuation rates were 0.68, 0.48, and 0.46 h^-1 for G. brevifissura, P. hessei, and Palaemon sp., respectively. The rate of gut pigment destruction for G. brevifissura was 99.6% of the total ingested, one of the highest values ever recorded for any crustacean. A gut pigment destruction of 79.0% was measured for Palaemon sp., 95.7% for P. hessei, and 93.8% for A. natalensis. During winter the total grazing impact of the dominant zooplankton species ranged from 5.05 to 22.7 mg chl-a m^-2 day^-1 and accounted for 34-69% of the available chl-a in the water column. During summer, the grazing impact ranged between 0.45 and 0.65 mg chl-a m^-2 day^-1, accounting for 17-41% of the available chl-a in the water column. This shows that the dominant zooplankton species of the Mpenjati have a very high grazing impact on algal cells. At times this may exceed 100% of the available phytoplankton production, suggesting that the zooplankton community may often resort to other food sources to meet all its energetic demands.     

Population dynamics and secondary production of the wedge clam <Emphasis Type="Italic">Donax</Emphasis><Emphasis Type="Italic">hanleyanus</Emphasis> (Bivalvia: Donacidae) on a high-energy,subtropical beach of Brazil

The population biology of Donax hanleyanus (Philippi, 1845) (Bivalvia: Donacidae) was studied by monthly sampling from June 1998 through June 2000, at Restinga da Marambaia Beach, Brazil. Two transects were established and divided into ten strata parallel to the waterline, and five replicates were taken with a 0.04 m² sampler in each stratum. The highest densities of D. hanleyanus were recorded in winter (September 1998, July 1999) and autumn (April 2000). A stratified distribution was observed: recruits were found mainly in the middle swash zone, while juveniles and adults occurred across the tidal gradient up to the retention zone. Mortality rates did not differ significantly between years. Annual production (in ash-free dry mass) ranged from 0.76 g AFDM m^-2 year^-1 (1998) to 3.67 g AFDM m^-2 year^-1 (1999), while the production-to-biomass (P/B) ratio varied from 1.45 to 1.59. Life span was ca. 17 months. Of all variables tested, only two, one biological and one physical, seem to have influenced the population dynamics of this species. A significant negative correlation between the densities of the suspension-feeders D. hanleyanus and Emerita brasiliensis (Crustacea: Decapoda) indicated possible intraguild competition, and there was a significant exponential correlation between beach slope and the mean across-shore position of D. hanleyanus. Data compiled from literature suggests a latitudinal gradient in population parameters, with mortality and renewal rate (P/B) of different species of Donax increasing and life span decreasing from temperate to tropical regions. Other factors influencing population dynamics, such as food availability, and the contributions of individual and community biomass to energy and nutrient cycling are also discussed.     

Differential effects of Aureococcus anophagefferens isolates ("brown tide") in unialgal and mixed suspensions on bivalve feeding

Picoplanktonic brown tides of Aureococcus anophagefferens have had devastating effects on production of commercially exploited bivalve populations in shallow, mid-Atlantic estuaries in the United States. The toxin produced by this alga has not been chemically characterized. This study develops a bioassay using juvenile mussels, Mytilus edulis, based on the inhibitory effect of brown tide on bivalve suspension-feeding, to compare the cellular toxicity of three Long Island, New York, clonal isolates of A. anophagefferens. Two recent (1995) isolates (CCMP 1707 and 1708) from Peconic Bay proved highly toxic and caused greater than 100-fold reduction in clearance rates (CR) of juvenile mussels in unialgal and mixed suspensions with a nutritious alga, Isochrysis galbana (clone T-iso), relative to controls. A third 1986 isolate from Great South Bay (CCMP 1784) showed no detectable toxicity in 24-h trials, and may have lost its initial potency over more than a decade of laboratory culture. Identification of a non-toxic strain provides a useful tool for future research. Cultures of the toxic isolate CCMP 1708 in late-stationary growth phase were significantly more toxic than those in early-exponential phase. The threshold concentration of toxic A. anophagefferens cells that inhibits clearance on co-occurring phytoplankton species was determined for juvenile (10-mm) hard clams, Mercenaria mercenaria. Relatively low concentrations (⣃᎒³ to 50᎒³ cells ml^-1) of isolate CCMP 1708 were sufficient to sharply reduce clam CR of I. galbana. Calculations based on these results suggest that, at peak historical densities of M. mercenaria in Great South Bay, removal of A. anophagefferens at low cell densities by suspension-feeding benthos could provide an effective top-down grazing control mechanism to prevent the initiation of brown tide in shallow, inner bays.     

Assimilation of dietary free amino acids,peptides and protein in post-larval Atlantic halibut (<Emphasis Type="Italic">Hippoglossus</Emphasis><Emphasis Type="Italic">hippoglossus</Emphasis>)

With the aim of improving the formulation of diets for early stages of marine fish, post-larval Hippoglossus hippoglossus (56 days after first feeding; 23 mg dry weight) were tube-fed liquid diets of radiolabelled L-(¹⁴C) free amino acids (FAA diet), peptides (PEPT diet) and protein (PROT diet), in order to compare the kinetics of transfer from gut into the liver and carcass. The test diets were prepared using equal concentrations of amino acids (AA) and fed as a single pulse (0.3 µl, 2.2 µg AA). The PROT diet consisted of methylated-¹⁴C-labelled (mainly lysine) bovine serum albumin (BSA), the PEPT diet was the PROT diet digested with trypsin and the FAA diet consisted of an amino acid mixture based on BSA composition with added ¹⁴C-lysine. The common labelling with ¹⁴C-lysine permitted a comparison of the three diets. Based on dissection and distribution of the tracer in each compartment, the FAA diet was almost completely absorbed 1 h after the tube-feeding; PEPT and PROT diets required more time before they were processed by the digestive tract. The FAA diet was absorbed about eight times as fast as protein and about six times as fast as peptides from the gut. A low retention of label in the liver for all diets at all sampling times suggests that the liver does not store the AA at these absorption rates. The FAA diet had a higher assimilation efficiency (about 90%) than the PEPT diet (about 12%) and PROT diet (about 32%). FAAs, thus, appear to be superior to protein and peptides as a dietary source of amino acids in post-larval Atlantic halibut.     

Significance of food type for growth of ephyrae Aurelia aurita (Scyphozoa)

We studied growth of newly released Aurelia aurita ephyra larvae fed five different food types, including a large-sized copepod, a phytoflagellate, and suspended POM (particulate organic matter) made from bivalve meat. Experiments were run at saturated food concentration in two different temperatures over 10 days. The effect of small differences in temperature was inconsistent and interacted with the effect of food type, which, in turn, was highly significant. A low average growth rate (4-9% day^-1) was shown when feeding on the large-sized copepod Calanus finmarchicus (80 µg AFDW individual^-1), in spite of an extremely high daily ration of up to 1500% of body AFDW. When feeding on the cryptophyte Rhodomonas baltica (ca. 8 µm cell diameter), the ephyrae showed an average growth rate over the 10 day experiment of 7-11%, but with a considerably higher growth rate during the first days. Suspended POM generated an average growth rate of 7-9% day^-1, whereas fresh bivalve meat, manually placed into the stomach of the ephyra, gave an average growth rate of 12-14% day^-1. Artemia nauplii (ca. 3 µg AFDW individual^-1), used as a general reference, resulted in higher growth rates than any of the other food types (17-31% day^-1). We conclude that A. aurita ephyrae can capture and feed on phytoplankton, large copepods, and POM; that phytoplankton might be of nutritive significance early in development; and that the high quantity of large-sized copepods ingested is inefficiently converted to growth during early development. POM is a potential food source because of the ability of the ephyrae to encounter and ingest it, although concentration, size distribution, and nutritional composition of natural POM probably constrain its effect on growth.     

Pools of chlorophyll and live planktonic diatoms in aphotic marine sediments

Chlorophyll a and numbers of live pelagic diatoms were recorded from sediment depth profiles at 11 stations in the oligotrophic Øresund, Denmark, in late-June. Extraction efficiency of chlorophyll a analysed fluorometrically did not differ significantly between paired samples of frozen-thawed and fresh sediment. The depth profiles of chlorophyll a could be explained by a diagenetic model involving two different chlorophyll pools: one reactive pool declining exponentially with core depth, and one non-reactive pool, of about 1 µg Chl ml^-1 wet sediment, being constant with depth. The number of live diatoms, quantified by the dilution-extinction method, and expressed in terms of most probable number (MPN), declined from an average of about 300,000 g^-1 in the surface sediment to zero values at a depth of 13 cm. The number of live cells was significantly correlated with the sediment chlorophyll a, and the profiles of live cells as well as reactive chlorophyll followed the same exponential decline with core depth, suggesting that the main source of chlorophyll in the sediment was live pelagic diatoms. Taxonomic composition of diatoms in the sediment, dominated by the pelagic genera Chaetoceros, Thalassiosira and Skeletonema, matched the species composition in the water column 3 months earlier during the spring bloom. Regular recordings of the phytoplankton community in the water column showed that only these specific bloom species could be the source of the sediment content of diatoms and chlorophyll a. Further, the ratios between live cells and chlorophyll a were similar in the sediment and in the spring bloom. A conservative estimate of depth-integrated pools of diatoms in the sediment suggested that about 44% of the total phytoplankton biomass during the spring bloom was still present as live cells in the sediment after 3 months. This indicates that the spring bloom input to the sediment is not degraded immediately by the benthic fauna.     

Reproductive cycles of three dominant Calanus species in Disko Bay, West Greenland

This study describes the annual reproductive cycles of the three dominant Calanus species, C. finmarchicus, C. glacialis and C. hyperboreus, in Disko Bay (West Greenland) in relation to seasonal phytoplankton development. Relative abundance of females, copepodite stage V (CV) and males, and the developmental stage of the female gonad were examined from plankton samples collected at weekly to monthly intervals from May 1996 to June 1997 with a WP2 net or a pump. During spring and summer, egg production rates were determined. Females of all three species were present year round. Maximum relative abundance was reached by C. hyperboreus females at the beginning of February, by C. glacialis in mid-February, and by C. finmarchicus in April. All three species reproduced successfully in Disko Bay. Their reproductive cycles were considerably different with respect to the timing of final gonad maturation and spawning, and hence in their relation to seasonal phytoplankton development. In all three species, early gonad development took place during winter, before living food became plentiful, suggesting that these processes were largely food independent. Final gonad maturation and spawning in C. finmarchicus was related to the phytoplankton concentration, reflecting that final gonad maturation processes are food dependent in this species. C. glacialis females matured and spawned prior to the spring bloom. Our results indicate that first internal lipid stores and later ice alga grazing supplied final gonad maturation and egg production. Maximum egg production rates of C. glacialis were found in spring and summer, when the chlorophyll a concentration was high. Mature female C. hyperboreus were found from February until mid-April, when the chlorophyll a concentration was still low. In this species, reproductive activity was decoupled from phytoplankton development, and final maturation processes and spawning were solely fuelled by internal energy stores.     

Factors determining the hatching success of Antarctic krill <Emphasis Type="Italic">Euphausia superba</Emphasis> embryo: lipid and fatty acid composition

The present study addresses the effect of maternal diet on hatching success and condition of embryos and larvae of Antarctic krill Euphausia superba. Lipid and fatty acid content and composition were determined in field and laboratory samples. Developmental stages analyzed in embryos included: multiple-cell, gastrula, and limb-bud stages. Larval stages analyzed included: nauplius I, nauplius II, and metanauplius. Laboratory-reared embryos were spawned by gravid females incubated under three feeding groups: (1) phytoplankton mixture, (2) phytoplankton mixture and minced clam, and (3) phytoplankton mixture, minced clam, and commercial larval food. Hatching success was highest in group 3 (100%), lowest in group 1 (0%), and highly variable in field samples (0–48%). Lipid decreased slightly in embryos during embryonic development, while large decreases in lipid were found during nauplius development. High levels of 18:2(n-6), 20:4(n-6), and 22:6(n-3) observed with group 3 samples coincided with high hatching success in krill embryos. The ratio of 22:6(n-3)/20:5(n-3) also correlated to hatching success of embryos. The fatty acid profile of embryos in group 3 was similar to that of the field-collected embryos, reflecting the contribution of the commercial larval food in the maternal diet. In our study, the maternal diet was found to influence the fatty acid composition of embryos and in turn affects the hatching success of krill. Specific polyunsaturated fatty acids appeared to play important roles in embryogenesis in krill.     

The importance of micro and macro morphological variation in the adaptation of a sublittoral demosponge to current extremes

Sponges are known to show morphological acclimation in response to habitat variation. However, previous studies have concentrated on only one aspect of morphological variation, either gross morphology or spicule morphology. Cliona celata (Grant) is common in a variety of different habitats on the south-west coast of Ireland and has been investigated with respect to morphological variability on both scales. C. celata exhibited six different gross morphological body forms (ridged, burrowing, massive, massive/chimneys, encrusting, encrusting/chimneys). The body form exhibited was correlated to local environment, showing the extent of morphological adaptation in C. celata. Sponge size varied (from 548ᇟ to 2,345녹 cm²) between sites, with the largest (2,345녹 cm²) being found at the most stable site where flow rates were <5 cm^-1 (F>23.24, P<0.05). This may seem paradoxical as growth conditions were considered poor, but mortality and damage from material in suspension was reduced at low energy sites. At the spicule level, morphological variation was also present. Spicules at high energy sites were significantly longer, narrower and less numerous than at low energy sites (F>15.36, P<0.05). Previously, spicule variation has been associated with increased stiffness in hostile environments. However, longer, thinner spicules, as found in C. celata, may result in a more flexible sponge. This is the first study to show both gross morphological (macro) and spicule (micro) variation in a single species of sponge. However, this study only reinforces some of the previously produced information on both of these adaptations of sponges to varying environments. This study also illustrates how the results of single studies should not be used to draw conclusions for group level adaptation.     

Energy budgets and feeding rates of Coryphaenoides acrolepis and C. armatus

This study develops energy budgets and estimates feeding rates for two macrourid fishes, Coryphaenoides acrolepis, dominant in the bathyal eastern North Pacific, and the abyssal cosmopolitan species, Coryphaenoides armatus. Daily energy expenditure by C. acrolepis was nearly twice that of C. armatus. C. acrolepis allocated nearly equal amounts of energy to metabolism and growth. Once sexual maturity was reached reproduction became the dominant energetic cost. Either these costs are necessary to retain adequate numbers of eggs and larvae on the continental slopes, or this fish does not reproduce on an annual basis and the calculated costs are an overestimate. C. armatus allocated relatively more energy to metabolism than growth. It may be semelparous, and this strategy would be of great energetic savings in its food-poor but stable environment. Individual daily ration for C. acrolepis decreased from 0.31% to 0.07% of body weight (BW) and for C. armatus from 0.12% to 0.02% BW with increasing fish length. These rates are substantially lower than those for fishes living in cold waters on the continental shelves. The population feeding rates for C. acrolepis ranged from 0.8 to 15 kg km^-2 day^-1 and for C. armatus from 5 to 2,800 g km^-2 day^-1. The scavenging behaviour of C. acrolepis was used to investigate the role of carrion as a food supply to the deep-sea benthos. It was estimated that the carrion eaten by C. acrolepis is equivalent to 0.04 mg C m^-2 day^-1 or only 0.2-0.4% of the average small particulate flux. Carrion consumption is important for scavengers like C. acrolepis, but it is not an important component of the carbon flux into the deep-sea benthic environment.     

Biomass, photosynthesis and growth of Laminaria saccharina in a high-arctic fjord, NE Greenland

Biomass, photosynthesis and growth of the large, perennial brown alga Laminaria saccharina (L.) Lamour. were examined along a depth gradient in a high-arctic fjord, Young Sound, NE Greenland (74°18'N; 20°14'W), in order to evaluate how well the species is adapted to the extreme climatic conditions. The area is covered by up to 1.6-m-thick ice during 10 months of the year, and bottom water temperature is <0°C all year round. L. saccharina occurred from 2.5 m depth to a lower depth limit of about 20 m receiving 0.7% of surface irradiance. Specimen density and biomass were low, likely, because of heavy ice scouring in shallow water and intensive feeding activity from walruses in deeper areas. The largest specimens were >4 m long and older than 4 years. In contrast to temperate stands of L. saccharina, old leaf blades (2-3 years old) remained attached to the new blades. The old tissues maintained their photosynthetic capacity thereby contributing importantly to algal carbon balance. The photosynthetic characteristics of new tissues reflected a high capacity for adaptation to different light regimes. At low light under ice, or in deep water, the chlorophyll a content and photosynthetic efficiency (!) were high, while light compensation (E_c) and saturation (E_k) points were low. An E_c of 2.0 µmol photons m^-2 s^-1 under ice allowed photosynthesis to almost balance, and sometimes exceed, respiratory costs during the period with thick ice cover but high surface irradiance, from April through July. Rates of respiration were lower than usually found for macroalgae. Annual elongation rates of leaf blades (70-90 cm) were only slightly lower than for temperate L. saccharina, but specific growth rates (0.48-0.58 year^-1) were substantially lower, because the old blades remained attached. L. saccharina comprised between 5% and 10% of total macroalgal biomass in the area, and the annual contribution to primary production was only between 0.1 and 1.6 g C m^-2 year^-1.     

Ostracod species-specific utilisation of sediment detritus and newly settled cyanobacteria, Aphanizomenon sp., in the Baltic Sea: evidence from stable carbon isotopes

The three meiobenthic ostracod species Candona neglecta, Paracyprideis fennica and Heterocyprideis sorbyana are commonly encountered in the deep (20-40 m) soft bottoms of the Baltic proper, and may contribute more to the total meiobenthic biomass than any other group. Experimental data indicate substantial differences in their utilisation of settling phytodetritus, with C. neglecta able to exploit newly settled organic material to a larger extent than P. fennica and H. sorbyana. Ostracod species composition, as well as ostracod and sediment carbon isotope contents, were studied in the field from an area with local differences in potential food resources. The study was performed during the late period of summer blooms of the cyanobacteria Aphanizomenon sp. Results showed that when all samples were taken together, adult C. neglecta (-22.4‰) was significantly more depleted in '¹³C than adult P. fennica (-21.0‰) and H. sorbyana (-20.3‰), indicating differences in food selection among the species. The flocculent sediment layer had, in all instances, lighter carbon ratios than did the lower layers. This trend was mirrored in most cases in the slight enrichment in C. neglecta compared to generally greater enrichment in the other two species. Carbon signatures of C. neglecta also varied significantly between stations, indicating that this species fed on different resources depending on location. Juvenile C. neglecta were far more depleted in '¹³C than adults and reflected the carbon signature of the cyanobacteria Aphanizomenon sp. The latter is known to be the most '¹³C-depleted phytoplankton member in the area.     

Studies on pigments,proteins and photosynthetic rates in some mangroves and mangrove associates from Bhitarkanika,Orissa

Pigment contents, proteins and net photosynthesis were investigated in fully developed leaf of 1-year-old seedlings of six mangroves (Bruguiera gymnorrhiza, Rhizophora apiculata) and mangrove associates (Caesalpinia bonduc, Cerbera manghas, Derris heterophylla, Thespesia populnea), collected from Bhitarkanika, located on the east coast of India. Large variations in the photosynthetic rates (P_N) among the six species were observed, ranging from 10.16 µmol CO₂ m^-2 s^-1 in C. bonduc to 15.28 µmol CO₂ m^-2 s^-1 in R. apiculata. The total leaf protein content ranged from 12.09 mg g^-1 dry wt in T. populnea to 51.89 mg g^-1 dry wt in B. gymnorrhiza. The chlorophyll a/b ratio was typically about 3.0 in all the studied species, except C. bonduc (2.8). Photosynthetic rates and chl a/b ratio in the leaves were found to be correlated. Analysis of chlorophyll and xanthophyll spectra suggested: (1) variations in different forms and amounts of carotenes as well as xanthophylls and (2) the presence of high amounts of near-UV-absorbing substances in leaves, particularly in the two mangroves (B. gymnorrhiza, R. apiculata) and a mangrove associate (T. populnea), which appears to be an adaptive feature. Estimation of the chl a/b ratios in isolated thylakoids yielded a low value of 1.8 for R. apiculata and >2.6 for other species. The total protein/chlorophyll ratios in thylakoids varied considerably from 3.14 (D. heterophylla) to 10.88 (T. populnea) among the mangrove associates and from 16.09 to 18.88 between the members of the Rhizophoraceae. The chlorophyll/carotenoid ratios in thylakoids of the six species were more or less similar. The absorption spectra for washed thylakoids of C. manghas and D. heterophylla exhibited absorption characteristics typical for C₃-plant thylakoids. However, thylakoids isolated from R. apiculata, B. gymnorrhiza, C. bonduc and T. populnea exhibited an unusual increase in absorption in the blue region (380-410 nm) of the absorption spectrum. The presence of high-absorbing (in the short-wavelength, near-UV region) pigments appears to be closely associated with the thylakoids in R. apiculata and T. populnea. Our results, therefore, suggest a wide range of variation, not only in protein and pigment contents of photosynthetic tissues, but also in the spectral characteristics and composition of the pigments in mangrove species. An understanding of the nature of these pigments in mangroves and their associates, under their natural conditions and especially in relation to eco-physiological adaptations, is necessary, not only in relation to conservation, but also to allow propagation under different salinity conditions.     

Estimating the influence of prawn stocking density and seagrass type on the growth of juvenile tiger prawns (Penaeus semisulcatus): results from field experiments in small enclosures

Fine mesh enclosures (0.9 m² in basal area, 1 m high, with 100 µm mesh) and a jet-net retrieval system were developed to test the influence of juvenile prawn stocking density on growth rates in (1) different months (April and October/November) and (2) different types of intertidal seagrass beds in the Embley River estuary of tropical Australia. Small juvenile tiger prawns (3-6 mm in carapace length, CL) were stocked in enclosures at densities of 4-32 prawns per enclosure (4.4-35.5 prawns m^-2) on a high biomass seagrass bed (about 70 g m^-2 of mostly Enhalus acoroides) and one with low biomass (about 10 g m^-2 of mostly Halodule uninervis). After 2-3 weeks in the enclosures, recovery rates, and hence possibly survival, were greater on the high biomass Enhalus than on the low biomass Halodule. However, not all fish and crustaceans could be excluded from the enclosures. Growth rates were twice as fast on the high biomass Enhalus than on the low biomass Halodule. It is likely that the high biomass Enhalus, with its greater surface area, supported more epiphytic flora and fauna and reduced the potential for interference competition between prawns, compared with the low biomass Halodule. Growth rates on Enhalus were significantly faster at a stocking density of 4 prawns per enclosure (1.3 mm CL week^-1) than at a stocking density of 16 and 32 prawns per enclosure (both 0.8 mm CL week^-1), and did not differ significantly between April and October/November (temperatures were about 30°C at both times). The mean growth rate at 8 prawns per enclosure (1.1 mm CL week^-1) did not differ significantly from those at 4, 16 and 32 prawns per enclosure. These results from two seagrass beds suggest that the carrying capacity for juvenile tiger prawns was greater in the high biomass Enhalus than the low biomass Halodule bed.     

Feeding biology and carbon budget of the sediment-burrowing heart urchin Brissopsis lyrifera (Echinoidea: Spatangoida)

This study of the burrowing heart urchin Brissopsis lyrifera includes measurements on feeding and food selection, ingestion rate, absorption efficiency, ventilation and respiration. B. lyrifera regulated feeding depth, ingestion rate and absorption efficiency in relation to food source. When food was added to the top layer of sediment, B. lyrifera burrowed closer to the surface and ingested mainly enriched surface material, whereas it burrowed deeper and ingested deep-seated sediment in the controls. In non-enriched sediment, the feeding rate was 0.04 g sediment DW h^-1 ind.^-1, and, in macro- and microalga-enriched sediment the feeding rate was 0.06 and 0.08 g sediment DW h^-1 ind.^-1, respectively. Absorption efficiency of TOC was about 43% in non-enriched sediment and in microalga-enriched sediment, but was significantly lower (34%) in macroalga-enriched sediment. Absorption efficiency of N varied between 48% and 55%, and was independent of food source. B. lyrifera feed selectively, enriching the gut TOC content about 2-fold and N content about 2.5-fold. The C/N ratio was therefore lower within the gut compared to the surrounding sediment, while the faecal C/N ratio was almost equal to the surrounding sediment. The faeces were, however, slightly richer in TOC and N compared to the surrounding sediment. For 3-5 cm long B. lyrifera, water current rate varied between 4 and 24 ml water h^-1, with a mean of 11 ml h^-1. Mean respiration rate was 205 µl O₂ h^-1 ind.^-1. The water current rate was not sufficient for B. lyrifera to sustain itself by filter feeding only. However, organic-rich particles from the surface are suggested to be an important contribution to the diet. A carbon budget was calculated for B. lyrifera from measured values of consumption, absorption efficiency and respiration, in order to estimate annual production of B. lyrifera. Compared to literature values, growth was overestimated about tenfold in the budget. A large proportion of the absorbed carbon was suggested to leave the animal as dissolved carbon, through mucus production or through anaerobic pathways, either by the heart urchin or by micro-organisms in the gut.