Occurrence of tetrodotoxin in the starfishes Astropecten polyacanthus and A. scoparius in the Seto Inland Sea

Two starfishes, togemomijigai Astropecten polyacanthus and momijigai A. scoparius were collected from the Seto Inland Sea in October 1983 through November 1984, and assayed for toxicity by the standard method for tetrodotoxin (TTX). Most of the 54 A. polyacanthus specimens assayed were toxic, with the highest toxicity score being 520 mouse units (MU) g^-1. All seven A. scoparius were toxic, with the highest score being 46 MU g^-1. The toxin from A. polyacanthus was purified by a method which consisted mainly of activated charcoal treatment, and chromatography on CM-Sephadex C-25 and Bio-Rex 70 columns. The purified starfish toxin showed a specific toxicity of 4 700 MU mg^-1, a value which was almost comparable to that of authentic TTX. From the thin-layer chromatographic and electrophoretic behavior, along with ¹H-NMR spectrum, A. polyacanthus toxin was identified as TTX.     

Rapana venosa as a bioindicator of environmental pollution

The aim of this investigation was to examine biochemical differences in the gastropod Rapana venosa (R. venosa) from polluted (RvP) and nonpolluted (RvN) sites of the Black Sea's Bulgarian coast that may serve as bioindicators of environmental quality. Mussels Mytilus galloprovincialis (MMg) were collected from polluted (MMgP) and nonpolluted (MMgN) sites at the same time for comparison. Bioactive compounds and the antioxidant activity of dry matter (DM) methanol extracts from RvP and RvN were determined. Three-dimensional fluorometry (3D-FL) and Fourier transform infrared (FTIR) analyses were used to evaluate polyphenols and proteins. DM methanol extracts from RvP, and to lesser extent RvN, showed high amounts of total phenolics (23.22±2.9 and 15.60±1.8 mg GAE · g^?1 DW) and exhibited high levels of antioxidant activity in some radical scavenging assays (μ MTE · g^?1 DW): 56.38±5.2 and 33.79±3.3 by 2,2azino-bis (3-ethyl-benzothiazoline-6-sulfonic acid) diamonium salt (ABTS·⁺); 54.79±5.6 and 33.7±2.6 by cupric reducing antioxidant capacity (CUPRAC); 62.34±6.8 and 30.31±3.9 by the 1diphenyl-2-picrylhydrazyl method (DPPH). A correlation in the above indices was found between R. venosa and M. galloprovincialis, but all results were relatively higher for M. galloprovincialis than for R. venosa. The obtained indices of R. venosa may serve as a bioindicator of environmental ecological quality.     

Consumption of diatoms and diatom filtrates by the tentaculate deposit-feederEupolymnia nebulosa (Annelida: Polychaeta)

Rates of organic uptakes of three live diatoms (Nitzschia acicularis, Nitzschia sp. andNavicula incerta), and of three corresponding filtrates by the deposit-feeding polychaeteEupolymnia nebulosa (Montagu) were measured under similar experimental conditions. Worms used during this study were collected by SCUBA diving at Port-Vendres in shallow water (7 m deep) during the summer of 1986. Uptake rates of live diatoms were affected both by length of the experiments and by the nature of the food offered. The highest rate of uptake (11.8 10^–4 mg algal ash-free dry wt mg^–1 worm dry wt h^–1) was recorded during a short-term experiment (4 h) with the smallest diatom (Nitzschia sp.). The lowest rate (1.1 10^–4 mg algal ash-free dry wt mg^–1 worm dry wt h^–1) was recorded during a long-term experiment (48 h) with the largest diatom (Nitzschia acicularis). Filtrates ofN. acicularis were more readily utilized than those ofNitzschia sp. andNavicula incerta. Because of differences in uptake rates of algal filtrates as a function of species, it is not possible to evaluate the bias due to interaction with dissolved substances in experimental studies assessing ingestion of live benthic diatoms byE. nebulosa.     

Feeding responses of the tentaculate deposit-feederEupolymnia nebulosa (Annelida: Polychaeta): Influence of sexual maturity

Feeding responses of the tentaculate depositfeeding polychaeteEupolymnia nebulosa (Montagu) were studied by measuring rates of uptake of three different¹⁴C-labelled diatoms (unialgal cultures ofNavicula incerta Grunow,Nitzschia acicularis Wm Smith, andNitzschia sp.). Worms used during this study were collected in the harbor of Port-Vendre (Western Mediterranean) during August 1986 (immature worms) and December 1987 (mature worms). Uptake rates were affected both by the length of the experiments and by the nature of the food offered. The highest rate of uptake (2.98 10^–4 mg ashfree dry wt of algae mg^–1 dry wt of worms h^–1) was obtained during short-term experiments (4 h) with the smallest diatom (Nitzschia sp.). The lowest rate of uptake (0.21 10^–4 mg ash-free dry wt of algae mg^–1 dry wt of worms h^–1) was also obtained withNitzschia sp., but for a long-term (48 h) experiment. There was no significant difference between rates of uptake of immature and mature worms.     

Resting egg formation and hatching of the S-type rotifer Brachionus plicatilis at varying salinities

Several live specimens of the blue-ringed octopus Octopus maculosus were collected from the Philippines in November 1985, and from Japan in February 1986, and the distribution of toxicity, along with toxin composition, in the posterior salivary gland and other soft parts were examined. Tetrodotoxin (TTX: 1400 mouse units g^-1) was detected in the posterior salivary gland of a Japanese specimen, while not only the salivary gland but other soft parts were toxic in the Philippine specimens. The Philippine specimens contained TTX and anhydrotetrodotoxin, the Japanese specimen TTX, 4-epitetrodotoxin, and an unknown toxin. The posterior salivary gland, intestine and other parts were excised from the Philippine specimens and examined for bacterial flora. Twenty-two dominant strains were isolated and cultured in a 2xORI medium (Ocean Research Institute, Simidu and Tsukamoto 1985) at 20°C for 20 to 48 h. Cells were harvested by centrifugation, and disrupted by ultrasonication. The toxins were partially purified from the cell lyzate by ultrafiltration and Bio-Gel P-2 column-chromatography. Instrumental analyses disclosed that 16 of the 22 strains produced TTX and/or related substances. Six strains which clearly exhibited TTX productivity were identified as Alteromonas (2 strains), Bacillus (2), Pseudomonas (1) and Vibrio (1), based on biochemical and biological characteristics. Of these, one strain each of Bacillus and Pseudomonas produced TTX at a level detectable by the mouse assay.     

Rapana venosa as an indicator species for TBT exposure over decadal and seasonal scales

Imposex and decline in reproductive output in marine gastropods have been linked to tributyltin (TBT) exposure. This study describes Chesapeake Bay, USA, veined rapa whelk Rapana venosa imposex incidence and sex ratios from 1998 to 2009. Tissue TBT concentrations (ng g^?1) were examined with respect to whelk sex, size, and water temperature at the time of collection, and also to egg case size, hatching success, and veliger diameter. Imposex incidence declined and population sex ratios moved closer to parity from 1998 to 2009. Observed TBT concentrations (ng g^?1) were higher in James River than in Ocean View whelks with higher TBT concentrations observed in males than in females. Exponential declines in TBT concentrations from female-specific first to last clutches within a reproductive season were observed, indicating that whelks depurate TBT through egg case deposition. Egg capsule hatching success and veliger size were similar for female and imposex whelks. The R. venosa imposex levels observed in Chesapeake Bay apparently do not affect the production, release, or viability of larvae.     

Diurnal gut pigment rhythm and metabolic rate of<Emphasis Type="Italic"> Calanus euxinus</Emphasis> in the Black Sea

The vertical distribution, diel gut pigment content and oxygen consumption of Calanus euxinus were studied in April and September 1995 in the Black Sea. Gut pigment content of C. euxinus females was associated with diel vertical migration of the individuals, and it varied with depth and time. Highest gut pigment content was observed during the nighttime, when females were in the chlorophyll a (chl a) rich surface waters, but significant feeding also occurred in the deep layer. Gut pigment content throughout the water column varied from 0.8 to 22.0 ng pigment female^–1 in April and from 0.2 to 21 ng pigment female^–1 in September 1995. From the diel vertical migration pattern, it was estimated that female C. euxinus spend 7.5 h day^–1 in April and 10.5 h day^–1 in September in the chl a rich surface waters. Daily consumption by female C. euxinus in chl a rich surface waters was estimated by taking into account the feeding duration and gut pigment concentrations. Daily carbon rations of female C. euxinus, derived from herbivorous feeding in the euphotic zone, ranged from 6% to 11% of their body carbon weight in April and from 15% to 35% in September. Oxygen consumption rates of female and copepodite stage V (CV) C. euxinus were measured at different temperatures and at different oxygen concentrations. Oxygen consumption rates at oxygen-saturated concentration ranged from an average of 0.67 g O₂ mg^–1 dry weight (DW) h^–1 at 5°C to 2.1 g O₂ mg^–1 DW h^–1 at 23°C for females, and ranged from 0.48 g O₂ mg^–1 DW h^–1 at 5°C to 1.5 g O₂ mg^–1 DW h^–1 at 23°C for CVs. The rate of oxygen consumption at 16°C varied from 0.62 g O₂ mg^–1 DW h^–1 at 0.65 mg O₂ l^–1 to 1.57 g O₂ mg^–1 DW h^–1 at 4.35 mg O₂ l^–1 for CVs, and from 0.74 g O₂ mg^–1 DW h^–1 at 0.57 mg O₂ l^–1 to 2.24 g O₂ mg^–1 DW h^–1 at 4.37 mg O₂ l^–1 for females. From the oxygen consumption rates, daily requirements for the routine metabolism of females were estimated, and our results indicate that the herbivorous daily ration was sufficient to meet the routine metabolic requirements of female C. euxinus in April and September in the Black Sea.Communicated by O. Kinne, Oldendorf/Luhe     

Distribution of tetrodotoxin in various organs of the starfish Astropecten polyacanthus

Tissues of various organs of the starfish Astropecten polyacanthus collected in the Hiroshima Prefecture from April 1985 to June 1986 were analyzed for lethal potency by the assay method for tetrodotoxin. The ovary showed the highest potency (47 to 1 450 MU g^-1), followed by the digestive organs (<10 to 960 MU g^-1) and the exoskeleton including spines and tube-feet (<10 to 170 MU g^-1). The pyloric caecum and testis were less toxic. Overall toxicity was remarkably higher in females (2 060±382 MU, mean±SE) than in males (1 106±214 MU).     

Mucous sheet formation on poritid corals: An evaluation of coral mucus as a nutrient source on reefs

The production, release, and subsequent consumption of coral mucus on reefs has been portrayed as a potential pathway for the transfer of coral and zooxanthellae production to other reef organisms. However, reported mucus production rates and analyses of nutritional value vary widely. Poritid corals provide a test system to measure mucus production because they produce mucous sheets which can be collected quantitatively. Fluid mucus and mucous sheets were collected fromPorites astreoides, P. furcata, P. divaricata during 1986 and 1987 on reefs in the San Blas Islands, Panama, La Parguera, Puerto Rico and the Florida Keys, USA. Mucus samples were collected from Indo-pacific poritids (P. australiensis, P. lutea, P. lobata, andP. murrayensis) on the Great Barrier Reef during 1985. Biochemical analyses of the fluid mucous secretions, and the derivative mucous sheet, indicate that the mucus is primarily a carbohydrateprotein complex.Porites fluid mucus had a mean caloric content of 4.7 cal mg^–1 ash-free dry weight (AFDW), while mucous sheets contained 3.5 cal mg^–1 AFDW. Sixty-eight percent of the mucous sheet was ash, while fluid mucus was 22% ash. The high ash and low organic contents suggest that mucous sheets have a low nutritional value. C:N ratios varied (range 6.9 to 13.7 for fluid mucus, and 4.8 to 5.9 for mucous sheets), but were similar to typical C:N ratios for marine organisms. Bacterial numbers and chlorophyll a concentrations were higher on mucous sheets than in the surrounding water. Although bacteria aggregate on mucous sheets, bacteria accounted for less than 0.1% of the carbon and nitrogen content of the mucous sheet. Lower C:N ratios in aged mucus, i.e. mucous sheets versus fluid mucus, were attributed to a loss of carbon rather than an increase in nitrogen. Mucous sheet production accounts for a small proportion (< 2% gross photosynthesis) of published values for coral production. In the San Blas Islands, Panama,P. astreoides produced mucous sheets at a rate of 1.5 g C m^–2 y^–1 and 0.3 g N m^–2 y^–1.P. astreoides andP. furcata produced mucous sheets with a lunar periodicity and may provide approximately monthly pulses of carbon and nitrogen to the reef food-web. However, the low annual production rates suggest that mucous sheets make a small contribution to overall energy flow on coral reefs.     

Algal chloroplasts in the protoplasm of three species of benthic foraminifera: taxonomic affinity,viability and persistence

The ultrastructure and pigment content of algal chloroplasts (derived from Bacillariophyceae or Chrysophyceae) are described from 3 benthic species of brackish-water foraminiferans.Elphidium williamsoni Haynes contains 4×10⁶ chloroplasts mg^-1, whereas the contents ofNonion germanicum (Ehrenberg) andE. excavatum (Terquem) are about 10% of this value. The two former contain chlorophyllsa andc and fucoxanthin, but these pigments were not detectable in the latter.E. williamsoni andN. germanicum had a net uptake of¹⁴C–HCO ₃ ^- , proportional to their content of chlorophyll and number of chloroplasts, increasing linearly up to approximately 10 Klux. At light saturation the former assimilates 2.3x10^-3 mg C mg^-1 h^-1 and the latter only about 20% of this value. Dark uptake was insignificant in all cases. Uptake could not be demonstrated inE. excavatum. The photosynthesis effected by these species is trivial in terms of the total benthic carbon fixation effected by the microflora. The chloroplasts survived longer in forminiferans kept in the dark than in light/dark adapted individuals. To keep a steady state population of chloroplasts under light/dark conditions,E. williamsoni must eat at least 65 chloroplasts individual^-1 h^-1, whereas the minimum consuption rate inN. germanicum is 20.     

Tetrodotoxin in sinking particles from coastal waters

The occurrence of tetrodotoxin (TTX) in marine sinking particles was investigated. Sinking particles were collected in 1991 using sediment traps in the coastal area of Aburatsubo Inlet, Japan. TTX and related substances were analyzed by tissue culture bioassay, high performance liquid chromatography (HPLC) and gas chromatography-mass spectrometry (GC-MS). TTX and related substances were detected from six samples. The concentrations were estimated to be ca. 200 to 1000 ng g^-1 by the HPLC analyses. To our knowledge, this is the first report on the occurrence of TTX in particles in an aquatic environment. The present results indicate that sinking particles are one of the sources of TTX in the marine environment and that these particles play a role in the toxification of marine organisms.     

Ontogenetic changes in digestive enzyme activity of the spiny lobster,<Emphasis Type="Italic"> Jasus edwardsii</Emphasis> (Decapoda; Palinuridae)

Digestive enzyme profiles of puerulus, post-puerulus, juvenile and adult stages of the spiny lobster Jasus edwardsii Hutton were determined in order to identify ontogenetic changes in digestive capabilities and assess capacity to use dietary components and to exploit diets to meet nutritional requirements. Juvenile and adult lobsters exhibited a diverse range of enzymes, suggesting they can exploit varied diets. Proteolytic activity of trypsin, chymotrypsin and carboxypeptidase A (1.86–3.70 U mg^–1 digestive gland protein) was significantly higher than carbohydrase activity of amylase, -glucosidase, cellulase and chitinase (0.0014–0.38 U mg^–1 digestive gland protein), thus showing that dietary protein was more important than carbohydrate and that the lobster is carnivorous. These conclusions are consistent with other studies that found spiny lobsters to feed predominantly on crabs, bivalves, ophiuroids and sponges. Lipase activity (0.371 U mg^–1 digestive gland protein) was also relatively high, thus showing the importance of dietary lipid. Total activities (units per digestive gland) of every enzyme assayed increased significantly with lobster carapace length. There were few significant ontogenetic trends in specific enzyme activity (U mg^–1 digestive gland protein). Amylase and laminarinase specific activity was significantly higher in small lobsters than large lobsters (regression analyses, respectively, F_(1,23)=9.84, P=0.005; F_(1,11)=19.65, P=0.001), suggesting that carbohydrates including laminarin are more important in the diet of small juveniles. Trypsin, amylase and lipase activities were detected in all non-feeding puerulus stages, suggesting that feeding is not a cue for digestive enzyme production in J. edwardsii. Significant variations in total and specific activities of amylase (F_(1,3)=14.2, P=0.00; F_(1,3)=14.2, P=0.00) and trypsin (F_(1,3)=8.8, P=0.00; F_(1,3)=21.41, P=0.00) and a declining trend in lipase total activity between non-feeding puerulus stages suggests that they may be hydrolysing endogenous energy reserves to sustain their onshore swimming activity prior to settlement. Profiles suggest carbohydrate and lipid are utilised first, followed by protein. Consistently high levels of lipase in all puerulus stages (0.24–0.7 U digestive gland^–1; P>0.05) confirm the importance of lipid as a major energy substrate.Communicated by G.F. Humphrey, Sydney     

Some properties of calcium-activated adenosine triphosphatase from the hermatypic coralGalaxea fascicularis

Samples of the hermatypic coralGalaxea fascicularis were collected between April 1987 and April 1990 from coral reefs off Singapore (103 °45E; 1 °13N). Ca²⁺-activated adenosine triphosphatase (ATPase) activity was detected in the plasma-membrane-enriched heavy microsomal fraction ofG. fascicularis. The high affinity component hadKm andVmax values of 0.0021 mM and 0.050 µmol Pi mg^–1 protein min^–1, respectively; corresponding values for the low affinity component were 0.15 mM and 0.85 µmol mg^–1 protein min^–1. The activity of the high affinity component was inhibited 80 and 50%, respectively, by the anticalmodulin drugs calmidazolium and chlorpromazine. The low affinity component of the Ca²⁺-ATPase may represent activities of alkaline phosphatase, Ca²⁺-ATPase from membranes of mitochondria and endoplasmic reticulum, or calmodulin-dissociated plasma membrane Ca²⁺-ATPase resulting from the removal of Ca²⁺ by EDTA during the isolation process. The high affinity Ca²⁺-ATPase is probably the enzyme responsible for Ca²⁺ extrusion from the cells ofG. fascicularis. The high and low affinity components of this Ca²⁺-ATPase could use ATP and ADP as substrates. Maximum activities of both components were registered at pH 7 and at 45°C. Ruthenium red, a specific inhibitor of Ca²⁺-ATPase, inhibited the activities of the high and low affinity Ca²⁺-ATPase by 100 and 60%, respectively. Inhibition of the activities of both components was also observed with sulphydryl reagents (PCMB and mersalyl). However, DCMU, diamox, dinitrophenol, iodoacetate, fluoride, cyanide, ouabain, oligomycin B and L-phenylalanine had no effect on the enzyme activities.     

Respiration rates of<Emphasis Type="Italic"> Beroe ovata</Emphasis> in the Black Sea

Metabolic rates of the ctenophore Beroe ovata within the length range from 0.4 mm (newly hatched larvae) to 60 mm were investigated. At 20° the respiration rates (Q, µg O₂ ind.^–1 h^–1) of individuals with wet weights (W, mg) less than or greater than 100 mg changed according to the equations Q=0.093W^0.62 and Q=0.016W^0.99, respectively. The weight-specific respiration rate of the juvenile ctenophores with wet body weights of 0.021–100 mg diminished approximately 20-fold (from 0.35 to 0.017 µg O₂ mg^–1 h^–1, respectively), but did not change within the range from 100 to 30,000 mg. The difference in the slope of the regression lines seems to be attributable to the ontogenetic changes in B. ovata metabolism. For the tested temperature range of 10–28°, the mean Q₁₀ coefficient was equal to 2.17±0.5. The basal metabolism of B. ovata narcotised by chloral hydrate was 4.5±0.9 times lower than total metabolism. Such a metabolic range is considered to be characteristic of aquatic invertebrates with high levels of locomotory activity.Communicated by O. Kinne, Oldendorf/Luhe     

Utilization of a fumigated sediment by two benthic deposit-feeders:Abra alba (Mollusca: Bivalvia) andEupolymnia nebulosa (Annelida: Polychaeta)

The present study tested the utilization of dead microbial biomass by two benthic deposit-feeders:Abra alba (Wood) (Mollusca: Bivalvia) andEupolymnia nebulosa (Montagu) (Annelida: Polychaeta). Clams were collected in the Canet lagoon during spring 1989. Worms were collected in the Port-Vendres harbour during spring 1989. The¹⁴C-labelled (glutamic acid, 24 h) sediment used during the study was sterilized with 1% chloroform, washed with sterile seawater, and dried (60°C; 48 h). This sterilisation procedure, called fumigation is the least harmful to the sediment (Novitsky 1986). Both clams and worms were incubated in the presence of the fumigated sediment for 5, 10, 20, and 50 h. At the end of each experiment we recorded the radioactivity in four compartments: (1) sediment, (2) dissolved organic matter (DOM), (3) CO₂, and (4) animals. The radioactivity of the sediment was subdivided into five fractions: (i) soluble in 2N HCl, (ii) soluble in hot 5% trichloroacetic acid (TCA), (iii) soluble in 1N NaOH, (iv) soluble in hot 6N HCl, (v) residual (after combustion in a Leco carbon analyser). In the first set of experiments, after 20 h of incubation, 5.4 and 4.7% of the total radioactivity was taken up by clams and worms, respectively. However, a model revealed that this uptake could have been correlated with the release of radiolabelled DOM (33% of total radioactivity during the first 5 h). In order to test this assumption, we used the same protocol with three additional washes of the fumigated sediment. This resulted in a significantly lower uptake by the clams (1.9% of the total radioactivity byt = 50 h), whereas the worms exhibited an uptake similar to that in the initial experiment (5.1% of total radioactivity byt = 50 h). These results underline the importance of considering interactions with DOM when applying radiotracer techniques to the study of benthic food chains. The average ingestion rates of fumigated sediment byA. alba andE. nebulosa were 5.2 10^–2 mg sediment dry wt mg^–1 clam h^–1 and 3.5 10^–2 mg sediment dry wt mg^–1 worm h^–1, respectively, which is comparable to previous data reported for other deposit-feeding bivalves and polychaetes feeding on natural sediment or detritus. The low radioactivity recorded for CO₂ together with the similarity of the changes in the partitioning of the radioactivity within the sediment between control experiments and experiments carried out in the presence of clams or worms suggest low assimilation efficiencies. Therefore, the present study supports the fact that dead microbial biomass does not constitute an important food source for benthic deposit-feeders.     

Abundances,growth rates,and production of tropical neritic copepods off Kingston,Jamaica

Weekly samples were collected near Kingston, Jamaica in 27 m vertical hauls, using 200 and 64µm mesh plankton nets, from July 1985 to January 1987. Thirtytwo copepod species were identified; nauplii and all copepodite stages were enumerated. Total copepod abundance ranged from 2.56 to 87.3 × 10⁴ m^–2. The annual abundance cycle was bimodal with peaks in October–November and May–June corresponding to the rainy seasons. Mean annual copepodite biomass was 0.15 g AFDW m^–2 ranging from 0.03 to 0.41 g AFDW m^–2. Mean generation time (from egg to adult) at 28°C was 19.5 d for the common speciesCentropages velificatus, Paracalanus aculeatus, andTemora turbinata. Isochronal development was demonstrated for copepodites ofP. aculeatus andT. turbinata, but not forC. velificatus. Mean daily specific growth rates (G) were 0.63, 0.63, and 0.48 d^–1 forC. velificatus, P. aculeatus, andT. turbinata, respectively. In general, daily specific growth rates decreased in the later copepodite stages. Thus, it is postulated that growth of later stages and egg production may be food limited. Annual copepodite production was estimated as 419 kJ m^–2 yr^–1, while annual exuvial production and naupliar production were 35 and 50 kJ m^–2 yr^–1, respectively. Egg production was estimated as 44% (184 kJ m^–2 yr^–1) of the total copepodite production. Thus, mean total annual copepod production was 688 kJ m^–2 yr^–1. This estimate is within the range of copepod production estimates in coastal temperate regions.     

Bacterial nitrification activity directly associated with isolated benthic marine animals

Potential nitrification rates (PNR) directly associated with isolated marine macrobenthic invertebrates were measured for a range of benthic epifaunal and infaunal species (bivalves, gastropods, polychaetes and crustaceans) collected from the Sacca di Goro, Po River delta, Italy. In the case of the filter-feeding bivalves, Tapes philippinarum and Mytilus galloprovicialis the PNR associated with the shell surfaces and dissected animal tissues (gills, siphons and residual tissue) were determined separately, in order to assess the distribution of the nitrifier populations. Significant PNR was found associated with all the tested macrofaunal species with activities ranging between 12 and 2,250 nmol ind.^–1 day^–1 and specific activities between 150 and 18,400 nmol g^–1 dry weight day^–1. However, no simple relationships were observed between PNR and the animals taxonomic or functional group, or with animal comportment (infaunal or epifaunal) or size class, indicating that more complex interactions may regulate the degree of colonisation of the animals by nitrifier populations. Incubations of shells alone and dissected tissues of the bivalves T. philippinarum and M. galloprovicialis demonstrated that approximately 50% of the total PNR activity was associated with the shell surfaces and 50% with the internal animal tissues, with the highest specific activities of 950 and 1,970 nmol g^–1 dry weight day^–1 determined for the gills of T. philippinarum and M. galloprovicialis, respectively. Thus, specific relationships may exist between the nitrifiers and their animal hosts. Overall, our data indicate that the macrofaunal stimulation of nitrification and/or coupled nitrification–denitrification observed in previous studies may not be solely due to the animals burrow walls serving as sites for nitrification, but also to the fact that the internal and external surfaces of the animals themselves are also colonised by nitrifying bacteria. Tentative calculations based on reported animal densities in the Sacca di Goro and the determined PNRs indicate that animal-associated nitrifier populations could contribute significantly to overall nitrification rates in situ, although further experiments are required to determine to what extent the potential rates measured in this study are realised under in situ conditions.Communicated by R. Cattaneo-Vietti, Genova     

Primary production of phytoplankton at a frontal zone located at the northern slope of the Dogger Bank (North Sea)

In July 1988 a survey was made in the Dogger Bank area of the North Sea. As a result of wind stress the area was found to be frequently well mixed. At the northerly slope a transition zone was observed between the stratified central North Sea and the well-mixed Dogger Bank area. Low nutrient concentrations were observed in surface waters; especially for nitrate (<0,1µM). High concentrations of phosphate (>0,5µM), nitrate (>1µM), ammonium (>2µM) and silicate (>2µM) only prevailed below the thermocline. Chlorophylla values were below 1µg l^–1 near the surface. Enhanced values (up to 4µg l^–1) were observed in the deeper layer at the transition zone and just below the thermocline at well-stratified locations. At the transition zone high specific C-fixation rates (up to 100 mg C mg^–1 chla d^–1) at the surface indicated the presence of enhanced productivity. The compensation depth for primary production was found to coincide with a specific C-fixation rate of 5 mg C mg^–1 chla d^–1. At greater depths, phytoplankton was only found where tidally induced vertical mixing allowed a regular exposure to higher light intensities. Storms resulted in a rapid redistribution of chlorophylla and enhancement of the C-fixation rate in the upper layer of the water column.Publication No. 10 of the project Applied Scientific Research Netherlands Institute for Sea Research (BEWON)     

Characteristic features of body composition and metabolism in some interzonal copepods

Protein, lipid, phosphorus, and organic carbon contents, as well as electron transport system (ETS) activity, lactatedehydrogenase activity, and gut evacuation rate, were measured in four interzonal species of Pacific copepods:Calanus australis, C. pacificus, Eucalanus inermis, andE. elongatus f.hyalinus, collected at the upwelling areas off Peru (8°S) and California (27°N), and in the middle of the North Pacific (30°N), from February to April 1987. The two Eucalanidae species —E. inermis andE. elongatus — have distinctive biochemical and elemental body composition and rates of main physiological processes. Relative protein, lipid, phosphorus, and organic carbon contents (µg mg^–1 wet weight) in these species were, respectively, ca. 1/7 to 1/10, 1/5 to 1/20, 1/5 to 1/10, and 1/5 those inCalanus spp. Likewise, oxygen uptake rate per unit of wet weight (based on ETS activity) inE. inermis andE. elongatus was 5 to 10% of that in calanids; a similar difference was found in phosphorus excretion rate. In addition, gut evacuation rates inE. inermis andE. elongatus were ca. one-fifth of those inCalanus spp. Based on these data, we considered the eucalanids as belonging to a distinctive physiological group, figuratively named jelly-body copepods. In contrast with calanids, active lactatedehydrogenase has been found in the bodies ofE. inermis andE. elongatus, apparently allowing them to survive for a long time in layers of extremely low oxygen content (<0.2 ml l^–1). The adaptive value of physiological features in these eucalanids and typical calanids is compared.     

Correlation between photosensitivity and downwelling irradiance in mesopelagic crustaceans

The current study determined behavioral and electrophysiological photosensitivities for three species of mesopelagic crustaceans: Pasiphaea multidentata Esmark, 1866 (Decapoda: Pasiphaeidae), Sergestes arcticus Kröyer, 1855 (Decapoda: Sergestidae), and Meganyctiphanes norvegica M. Sars, 1857 (Euphausiacea: Euphausiidae). In addition, in situ quantifications of the species vertical distributions in relation to downwelling irradiances were also determined in two locations in the northwest Atlantic Ocean, Wilkinson Basin (WB) and Oceanographer Canyon (OC). Data are from six 2-week cruises between June and September from 1995 to 2001. P. multidentata and M. norvegica were the most abundant large crustaceans in WB, and S. arcticus and M. norvegica were the most abundant large crustaceans in OC. The behavioral light sensitivity thresholds of P. multidentata and M. norvegica from WB were both 10⁷ photons cm^–2 s^–1 and those of S. arcticus and M. norvegica from OC were both 10⁸ photons cm^–2 s^–1. Electrophysiologically, P. multidentata was significantly more sensitive than M. norvegica from either location, S. arcticus was significantly more sensitive than M. norvegica from OC, and M. norvegica from WB was significantly more sensitive than M. norvegica from OC. A correlation was found between electrophysiologically measured photosensitivity and downwelling irradiance, with the most sensitive species, P. multidentata and S. arcticus, associated with the lowest irradiance at daytime depths. The photosensitivities of M. norvegica collected from the clearer waters of OC were significantly lower than those of individuals collected from the more turbid WB waters. These results indicate that downwelling irradiance has a significant impact on interspecies and intraspecies vertical distribution patterns in the mesopelagic realm.Communicated by J.P. Grassle, New Brunswick