Ecological investigations of the zooplankton community of balsfjorden,northern norway: The genital system in Calanus finmarchicus and the role of gonad development in overwintering strategy

The structure of the reproductive system in Copepodite Stages IV and V of the marine copepod Calanus finmarchicus (Gunnerus) is described, together with seasonal variations in gonad development and sex ratio, from Balsfjorden (69°21N; 19°06E), a subarctic fjord in northern Norway. The genital system in Copepodite Stage IV consists of a small gonad with two genital ducts. The gonad of Copepodite Stage V in its most immature condition resembles that seen in Copepodite Stage IV, whereas the sex of Stage V copepodites with mature gonads can be easily and reliably determined. The genital ducts are the key characters for separating males and females in Copepodite Stage V; potential males have one and potential females two genital ducts connecting the gonad to the genital opening on the first urosome segment. Adults males were rare compared to females, and were only in the majority at the end of January and beginning of February. The sex-ratio also favoured females in Copepodite Stage V. Seasonal peaks in the proportion of adult males reflected peaks of males in Copepodite Stage V, suggesting that seasonal variations in the adult sex-ratio are a direct result of a situation which has already been determined in Copepodite Stage V. The seasonal variation in gonad development in Copepodite Stages IV and V of C. finmarchicus reveals that the size of maturity of the gonad varies throughout the year. This study indicates that visible sex-differentiation in Stage V and moulting into adults occurs in Balsfjorden at least 2 months before spawning in April. Phytoplankton levels are immeasurable before the spring diatom increase starts at the end of March, implying that development and maturation of gonads are dependent upon internal energy resources, whereas the final act of spawning seems to require energy input from phytoplankton.     

Respiration and excretion rates of Calanus glacialis in arctic waters of the Barents Sea

The respiration and excretion rates of Calanus glacialis (Jaschnov) Copepodite Stages III, IV, V, and adult females from the drift-ice area east of Svalbard (Barents Sea) were measured in shipboard experiments during the period from 27 May to 13 June, 1983. The phytoplankton biomass and abundance varied considerably between localities, but these variations were not generally reflected in the respiration and excretion rates of the copepod. The respiration and excretion rates of C. glacialis at the ambient temperature of-1.8°C (average respiration rates of 0.95, 0.73, 0.57, and 0.60 l O₂ mg^-1 dry wt h^-1 for Copepodite Stage III, IV, V, and adult females, respectively) were similar to those previously reported for other large-sized copepods from cold or temperate areas. Average respiration and excretion rates tended to decrease with incubation time or time after capture. Measurements on ten occasions within a period of 27 h after capture revealed excretion rates of ammonium ranging between 2.9 and 16.8 for C III, 3.7 and 21.1 for C IV, 1.3 and 28.4 for C V, and 1.6 and 18.7 for adult females, all expressed as nmol mg^-1 dry wt h^-1. In all experiments, excretion rates of inorganic phosphate varied between 0.7 and 1.5 (C III), 0.5 and 1.1 (C IV), 0.2 and 0.8 (C V), and 0.3 and 1.0 (adult females) nmol mg^-1 dry wt h^-1. Ratios of O:N, O:P, and N:P indicated that much of the metabolic energy was derived from catabolism of proteins. Comparison of the turnover rate of carbon and nitrogen showed, however, that nitrogen turnover was between 2.6 and 8.9 times higher than that of carbon. This may indicate that the copepods deaminate ingested protein, with the carbon skeleton of the amino acids subsequently being used in the synthesis of lipid compounds, possibly wax esters.     

Copepod grazing during a declining spring phytoplankton bloom in the Northern North Sea

Grazing rates of larger (Calanus finmarchicus) and smaller (Acartia clausii Pseudocalanus elongatus etc.) copepods on naturally occurring phytoplankton populations were measured during a declining spring phytoplankton bloom. During the initial period, dominated by Chaetoceros spp. diatoms, constant ingestion rates were observed in Calanus finmarchicus at suspended particulate concentrations above 300 g carbon l^-1. Average daily intake during this time amounted to 35 to 40% of body carbon and reached a maximum of 50%. The feeding response of the smaller copepods was not so well defined, although a maximum daily intake of 56% body carbon was recorded. In both groups, feeding thresholds were at particulate concentrations around 50 g C l^-1. The feeding response of C. finmarchicus was correlated with both a change in their own population and in the food cell type. Linear regressions describing the concentration-dependent feeding response were: ingestion rate (IR)=1.16 total particulate volume (TPV)-36.15 during the initial part of the period compared with IR=0.41 TPV-12.18 for the latter period. C. finmarchicus filtered out slightly larger (x 1.2 diameter) particles than the small copepods and, in both groups, some filtering adjustment was made to accomodate to modal changes in the phytoplankton population from 20–30 m to 10 m diameter cells. Particle production during feeding was frequently evident in the smallest size ranges of particles and the ratio of particle production to ingestion rate was greater at low feeding rates.     

Temperature and plankton

The relationship between temperature and metabolism was studied in Artic copepods with regard to the concept of metabolic cold adaptation of polar poikilotherms. Temperature tolerance and respiration rates of the dominant copepods Calanus finmarchicus (Gunnerus), C. glacialis (Jaschnov), C. hyperboreus (Krøyer) and Metridia longa (Lubbeck), collected in Fram Strait, Greenland Sea, in July 1983, were studied at different temperatures. Temperature tolerance in the boreal C. finmarchicus was slightly higher than in the three Arctic species. Respiration rates at lower temperatures followed the Arrhenius equation in all species, with values for (temperature characteristics) between 11.05 and 22.95, corresponding to a Q₁₀ between 2.05 and 4.5. This increase in metabolic rate with rising temperature was not related to an increase of swimming activity, as was shown by videoanalysis. Activity was determined as average swimming speed and as frequency of certain locomotor patterns. Average swimming speed remained unchanged at all temperatures and was ca 1 cm s^-1 for all species, when only periods of active swimming were considered. The time spent with active swimming did not change with temperature in M. longa and C. finmarchicus, but decreased in c. glacialis. In C. hyperboreus it increased at 5°C and decreased again at higher temperatures. It is suggested that the increase in oxygen consumption is fully accounted for by the basal metabolism.     

Comparative observations on the biometry and development of Calanus finmarchicus and C. helgolandicus in Copepodite Stage V,with comments on other Calanidae

A similar, sexual dimorphism in Copepodite Stage V was observed in winter generations of Calanus finmarchicus (Gunnerus) and C. helgolandicus (Claus), co-existing in the Firth of Clyde, Scotland. In both species, the frequency distribution of prosome length in the Copepodite V population was bimodal. Among individual Stage V copepodites kept in the laboratory, those of the larger form alone produced adult males. Indications of morphological distinctions between the forms other than in size, however, were absent. Differences in least-squares regressions of individual cephalosome on metasome lengths, previously attributed to a distinction in prosome shape by Grigg et al. (1985), reflected size alone. These two measurements also were greater in the “large” form, but this independently of each other. Variations in their relative contributions to the prosome length distinguished individuals, and were continuous between the forms. Present and previously published observations on members of the family Calanidae have been compared. The presence, or not, of a sexual distinction in prosome length in the Stage V copepodite is variable among calanids, within and between species. A possible association with the extent of seasonal variations in size and life-cycle characteristics is suggested.     

Egg production of the Arctic copepod Calanus glacialis: laboratory experiments

Egg production of single female Calanus glacialis Jaschnov fed with Thalassiosira antarctica at super-abundant concentrations (>300 g C l^-1) was determined over several weeks. Experiments were performed directly after collection from the East Greeland Current in June 1987 and 1988, and during resumed feeding after long-term starvation over 4 (October 1988), 4.5 (October 1987) and 6.5 (January 1988) mo. In addition, in June 1987, short-term starvation experiments of 5 and 15 d were conducted. Egg production was closely related to feeding in all experiments. While directly after collection eggs were produced within a few days, it took 2 wk (October 1987 and 1988) and 10 d (January 1988), respectively, to resume egg production after long-term starvation. During long-term starvation periods eggs were not laid. The decrease in total egg production with duration of starvation was due to decreasing clutch size and increasing spawning interval. In contrast, short-term starvation experiments only affected spawning interval. Interannual variability in egg production was high, with much higher clutch sizes in 1988. Average production rates in June 1988 correponded to 5% body C female^-1 d^-1, the maximum was 7.4% (1 274 eggs in 23 d). Carbon content of eggs was 0.40 g egg^-1. C. glacialis is well adapted to pulsed food events in the Arctic by its longevity; its ability to preserve its reproductive potential over several months; its rapid mobilization of ovaries; and by its high egg production rates. The implication of prolonged spawning capacity on life cycle studies is discussed.     

Feeding,growth and bioluminescence of the heterotrophic dinoflagellate Protoperidinium huberi

Feeding, growth and bioluminescence of the thecate heterotrophic dinoflagellate Protoperidinium huberi were measured as a function of food concentration for laboratory cultures grown on the diatom Ditylum brightwellii. Ingestion of food increased with food concentration. Maximum ingestion rates were measured at food concentrations of 600 g C l^-1 and were 0.7 g C individual^-1 h^-1 (1.8 D. brightwelli cells individual^-1 h^-1). Clearance rates decreased asymptotically with increasing food concentration. Maximum clearance rates at low food concentration were ca. 23 l ind^-1 h^-1, which corresponds to a volume-specific clearance rate of 5.9x10⁵ h^-1. Cell size of P huberi was highly variable, with a mean diameter of 42 m, but no clear relationship between cell size and food concentration was evident. Specific growth rates increased with food concentration until maximum growth rates of 0.7 d^-1 were reached at a food concentration of 400 g C l^-1 (1000 cells ml^-1). Food concentrations as low as 10 g C l^-1 of D. brightwellii (25 cells ml^-1) were able to support growth of P. huberi. The bioluminescence of P. huberi varied with its nutritional condition and growth rate. Cells held without food lost their bioluminescence capacity in a matter of days. P. huberi raised at different food concentrations showed increased bioluminescence capacity, up to food concentration that supported maximum growth rates. The bioluminescence of P. huberi varied over a diel cycle, and these rhythmic changes persisted during 48 h of continuous darkness, indicating that the rhythm was under endogenous control.     

Seasonal observations on the biometry and development in Copepodite Stage V of Calanus finmarchicus from the Firth of Clyde

Relationships between linear dimensions of casts produced at the final moult in the laboratory and definitive sex confirm that Calanus finmarchicus (Gunn.) is sexually dimorphic in Copepodite Stage V. Two forms of Stage V copepodite were present at all times of the year in the Firth of Clyde, Scotland. Except in early spring, when size was similar, these were consistently distinguished by the prosome length. The small form moulted to adult females and the large form to adults of either sex. Relative to the metasome, the length of the cephalosome was greater in the large than small form of Stage V copepodite. This, to an increased extent, distinguished males from all females in the adult. Differences between females of the large and those of the small form were similar in the Stage V copepodite and adult. Feminization in the large form entailed an arrest in the differentiation of secondary male sex-characters. Most morphological females moulting from both forms of Stage V copepodite possessed an apparently normal ovary, with eggs extending into oviducal diverticula. In the case of the large form, however, eggs were occasionally absent from diverticula. The gonad in two specimens resembled an immature testis. In males, a normal testis was always present. Differentiation into the adult male entailed the internal suppression of feminizing factors. On average, within collections, females of the large form moulted later than the small form and definitive males. A similarly consistent difference between females of the small form and males was absent. Moulting times in the large form of Stage V copepodite depended on the presence or not of feminization. Differentiation into the adult male required the environmental stimulation of development. Comparisons with moulting patterns in the sea suggest a mechanism that regulates the recruitment of males to the adult.     

Production of DMSP and DMS during a mesocosm study of an Emiliania huxleyi bloom: influence of bacteria and Calanus finmarchicus grazing

We investigated the influence of bacteria and metazooplankton on the production of dimethylsulfoniopropionate (DMSP) and dimethylsulfide (DMS) during blooms of Emiliania huxleyi (Lohmann) Hay and Mohler in seawater mesocosms. The phytoplankton succession was marked by the rapid collapse of an initial Skeletonema costatum (Greville) Cleve bloom followed by a small E. huxleyi bloom. The collapse of the diatom bloom was accompanied by an increase in concentrations of dissolved DMSP (DMSP_d) and bacterial abundance and activity (as determined by the thymidine incorporation technique). The increase in bacterial activity was followed by a rapid decrease in DMSP_d concentrations which remained low for the rest of the experiment, even during the subsequent collapse of the E. huxleyi blooms. The absence of DMSP_d and DMS peaks during the declining phase of the E. huxleyi blooms was attributed to the high bacterial activity prevailing at that time. The influence of metazooplankton grazing on DMSP and DMS production was investigated by adding moderate (24 mg dry weight m^-3) and high (520 mg dry weight m^-3) concentrations of Copepodite Stage V and adults of Calanus finmarchicus to two of four filtered (200 m mesh net) enclosures during the E. huxleyi blooms. The addition of C. finmarchicus, even in high concentrations, had no apparent effect on the dynamics of E. huxleyi, suggesting that the copepods were not grazing significantly on nanophytoplankton. The addition of copepods in high concentrations favored an accumulation of chlorophyll a and particulate DMSP. These results suggest that copepods were preying on the herbivorous microzooplankton which, in turn, was controlling the biomass of nanophytoplankton. DMS production was also enhanced in the enclosure with maximum metazooplankton biomass, suggesting that the grazing of C. finmarchicus on microzooplankton containing DMSP may contribute to DMS production. These results provide strong support to the emerging idea that bacteria and metazooplankton grazing play a dominant role in determining the timing and magnitude of DMS pulses following phytoplankton blooms.     

Nutrition studies in the nauplius larva of Calanus pacificus (Copepoda: Calanoida)

Nauplii of Calanus pacificus were raised on a mixture of algae. Details of the mouth-parts, such as denticles, labial palps and lobes, setations and structure of the masticatory teeth were examined by scanning electron microscopy (SEM). Under the experimental conditions (15°C and 300 gC l^-1), exponential growth coefficients for the period Nauplii II–VI were 0.179 for carbon and 0.228 for nitrogen. C:N ratios dropping from 5.1 to 4.7. Growth was isochronous, each stage lasting 1.5 days. Respiratory losses were 15 to 19.6% of body carbon daily. Nauplii raised on a given alga showed higher rates of ingestion in the presence of this food, compared to nauplii switched to other algae. Minimal threshold concentrations for feeding were found, depending on the size of the food offered and ranging from 5.8 gC I^-1 for Lauderia borealis (28.7 m spherical diameter) to 47.1 gC 1^-1 for Chlamydomonas sp. (11.0 m). Unlike the Copepodite I stage, Nauplii II–VI larvae were not able to ingest small cells such as Isochrysis galbana (4.3 m), or very large ones such as Ditylum brightwellii (47.5 m) at more than maintenance rations. Below the critical concentration for maximal feeding, ingestion was clearly dependent on size of the cells offered, but the size-dependent relationship was different for diatoms and non-diatoms. Filtering rates increased from a threshold concentration to a maximal rate at about 50 gC 1^-1, and decreased at higher concentrations. Critical concentrations ranged from 125 gC 1^-1 for L. borealis to 1000 gC 1^-1 for Chlamydomonas sp. Maximal daily rations ranged between 100 and 150% of body carbon.     

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     

Uptake of dissolved organic matter by larval stage of the crown-of-thorns starfish Acanthaster planci

The life-history of the crown-of thorns starfish (Acanthaster planci) includes a planktotrophic larva that is capable of feeding on particulate food. It has been proposed, however, that particulate food (e.g. microalgae) is scarce in tropical water columns relative to the nutritional requirements of the larvae of A. planci, and that periodic shortages of food play an important role in the biology of this species. It has also been proposed that non-particulate sources of nutrition (e.g. dissolved organic matter, DOM) may fuel part of the nutritional requirements of the larval development of A. planci as well. The present study addresses the ability of A. planci larvae to take up several DOM species and compares rates of DOM uptake to the energy requirements of the larvae. Substrates transported in this study have been previously reported to be transported by larval asteroids from temperate and antarctic waters. Transport rates (per larval A. planci) increased steadily during larval development and some substrates had among the highest mass-specific transport rates ever reported for invertebrate larvae. Maximum transport rates (J _max ⁱⁿ) for alanine increased from 15.5 pmol larva^–1 h^–1 (13.2 pmol g^–1 h^–1) for gastrulas (J _max ⁱⁿ=38.7 pmol larva^–1 h^–1 or 47.4 pmol g^–1 h^–1) to 35.0 pmol larva^–1 h^–1 (13.1 pmol g^–1 h^–1) for early brachiolaria (J _max ⁱⁿ just prior to settlement=350.0 pmol larva^–1 h^–1 or 161.1 pmol g^–1 h^–1) at 1 M substrate concentrations. The instantaneous metabolic demand for substrates by gastrula, bipinnaria and brachiolaria stage larvae could be completely satisfied by alanine concentrations of 11, 1.6 and 0.8 M, respectively. Similar rates were measured in this study for the essential amino acid leucine, with rates increasing from 11.0 pmol larva^–1 h^–1 (or 9.4 pmol g^–1 h^–1) for gastrulas (J _max ⁱⁿ=110.5 pmol larva^–1 h^–1 or 94.4 pmol g^–1 h^–1) to 34.0 pmol larva^–1 h^–1 (or 13.0 pmol g^–1 h^–1) for late brachiolaria (J _max ⁱⁿ=288.9 pmol larva^–1 h^–1 or 110.3 pmol g^–1 h^–1) at 1 M substrate concentrations. The essential amino acid histidine was transported at lower rates (1.6 pmol g^–1 h^–1 at 1 M for late brachiolaria). Calculation of the energy contribution of the transported species revealed that larvae of A. planci can potentially satisfy 0.6, 18.7, 29.9 and 3.3% of their total energy requirements (instantaneous energy demand plus energy added to larvae as biomass) during embryonic and larval development from external concentrations of 1 M of glucose, alanine, leucine and histidine, respectively. These data demonstrate that a relatively minor component of the DOM pool in seawater (dissolved free amino acids, DFAA) can potentially provide significant amounts of energy for the growth and development of A. planci during larval development.     

Uptake of glucose by bacteria in the sediment

A method is described for the incubation of undisturbed sediment cores under in situ conditions with the addition of low concentrations of ¹⁴C-glucose. Data are presented for respiration, gross uptake and actual uptake rate of glucose by bacteria in sandy, wave-washed beaches of the Baltic Sea. On the average, the bacteria respired 8% of the total glucose taken up. The gross uptake measured was between 2.3×10^-3 and 6.8×10^-3 g ¹⁴C-glucose g sediment^-1 (dry weight) h^-1 (average 4.7×10^-3 g g^-1 h^-1). Minima in the gross uptake rate corresponded with maxima in the concentration of natural free dissolved glucose. For the actual uptake rate, however, very similar uptake rates were calculated for the sediments examined (between 1.4×10^-1 and 1.9×10^-1 g glucose g^-1 h^-1, average 1.7×10^-1 g g^-1 h^-1).Publication No. 183 of the Joint Research Program at Kiel University (Sonderforschungsbereich 95 der Deutschen Forschungsgemeinschaft).     

Chemical composition,metabolic rates and feeding behavior of the midwater ctenophore Bathocyroe fosteri

Individuals of the midwater ctenophore Bathocyroe fosteri (0.01 to 1.6 g dry weight, DW) were collected from Bahamian waters by the submersible Johnson-Sea-Link during May and September/October 1983 and October/November 1984 from 530 to 700 m depth. Metabolic rates were measured and showed oxygen consumption to be in the range of 0.01 to 0.18 mg O₂ g^-1 DW h^-1 at temperatures ranging from 9° to 12°C. Ammonium excretion (0.01 to 0.14 g-at N g^-1 DW h^-1) was typically low. Energy expenditures estimated from respiration data (ca. 7% body C d^-1) indicated that one to three midwater crustacean prey (ca. 150 g C d^-1) could provide the daily maintenance ration required by a 40 mm ctenophore. These metabolic characteristics complemented in situ observations of poor locomotor ability and passive feeding behavior.     

Grazing,defecation and excretion rates of copepods from inter-island channels of the Canadian Arctic archipelago

Zooplankton (>800 m) and water samples were collected at night at eleven stations in Parry Channel and adjacent waters, between 28 August and 14 September 1986. Chlorophyll concentrations varied between 17 g l^-1 at the surface at one station in Wellington Channel, Canada, and 1.5 g l^-1 throughout the top 30 m at one station in Byam Martin Channel, Canada. In tows from 0 to 50 m the zooplankton community at all stations was dominated by varying proportions of three species,Calanus hyperboreus, C. glacialis andMetridia longa. Levels of chlorophyllderived pigments inC. hyperboreus Stages V, IV and III were correlated with the concentration of chlorophyll at the chlorophyll maximum. Defecation rate constants, measured for the different stages where present, were variable but not correlated with ambient chlorophyll concentrations. Ammonia excretion was measured simultaneously for the communities in which defecation was being measured in the invividual species and stages. In these experiments the rate of ammonia accumulation decreased significantly over the period during which the copepods were actively defecating (usually the first 3 h) and then tended to a constant level (over the next 18 to 20 h). The time courses of ammonia accumulation could be described by a model comprised of the sum of a straight line and a saturating curve. For seven experiments the ammonia release given by the asymptote of the saturating component was correlated with the estimate of community defecation, obtained by summing the individual defecations, suggesting that the two processes were closely associated. Weight specific defecation and weight specific defecation-associated ammonia excretion were both correlated with ambient chlorophyll concentration. The ratio of initial to basal ammonia excretion rate varied between 2 and 20, so that defecation-associated ammonia release may be important in the estimation of in vivo nitrogen excretion or regeneration rates.     

Ammonium excretion by gelationous zooplankton and their contribution to the ammonium requirements of microplankton in Chesapeake Bay

Ammonium excretion rates of recently collected specimens of gelatinous zooplankton, the scyphomedusan Chrysaora quinquecirrha DeSor and the etenophore Mnemiopsis leidyi A. Agassiz, were correlated with body mass and water temperature in measurements made from April to October 1989 and 1990. Rates ranged between 3.5 and 5.0 g atoms NH ₄ ⁺ -N (g dry wt)^-1h^-1 for C. quinquecirrha and 3.0 to 4.9 g atoms NH ₄ ⁺ -N (g dry wt)^-1h^-1 for M. leidyi. Excretion rate equations and in situ data on the size distributions and biomasses of gelatinous zooplankters and water temperature were used to estimate the contribution of ammonium by medusae and ctenophores to mesohaline Chesapeake Bay waters on several dates during April to October 1989 and 1990. We then compared the estimated contributions to direct measurements of ¹⁵NH ₄ ⁺ uptake by microplankton. The maximum estimated regeneration by gelatinous zooplankton was 5.8 g atoms NH ₄ ⁺ -N m^-3h^-1 at night in August 1990, when medusae biomass was greatest. This represents about 4% of the ammonium required by the microplankton. During the daytime on all dates, less than 1% of the ammonium required by microplanktion was supplied by gelatinous zooplankton. Therefore, gelatinous zooplankton appear to play a minor role in the ammonium cycle of Chesapeake Bay.     

Calcification in the maerl coralline alga Phymatolithon calcareum: Effects of salinity and temperature

The coralline alga Phymatolithon calcareum was dredged from 13 m in the Kattegatt, Baltic Sea, in December, 1980, and its rate of calcification was measured by ⁴⁵Ca⁺⁺-uptake methods. Light-saturated calcification rates at 5°C ranged from 15.8 g CaCO₃ g^-1 dry wt h^-1 for the basal parts of the plants to 38.7 g CaCO₃ g^-1 dry wt h^-1 for the tips. These age gradients were not apparent when calcification rates were expressed on the basis of surface area. Experiments with salinity (10, 20, 30) and temperature (0°, 5°, 10°, 20°C) indicated that optimum conditions for calcification were at 30 S and at temperatures above 10°C. Salinity had a greater influence on calcification rate than did temperature, and there was a positive relationship between salinity and calcification rate at all temperatures. In 6 mo old cultures, salinity was again the important factor, with all plants remaining healthy at 30 except those at the highest temperature (20°C). These trends, and the low calcification rates at 10S (4.6 g CaCO₃ g^-1 dry wt h^-1 at 5°C to 8.6 g CaCO₃g^-1 dry wt h^-1 at 20°C) suggest that low salinity may be the explanation for the general absence of P. calcareum from the brackish waters of the Baltic Sea. Short-term experiments in which salinity was kept constant while Ca⁺⁺ concentration was altered, and experiments in which salinity was varied and Ca⁺⁺ concentration kept constant, suggest that it is the calcium ion concentration and not salinity per se which affects calcification rates.     

The common jellyfish Aurelia aurita: standing stock,excretion and nutrient regeneration in the Kiel Bight,Western Baltic

The population dynamics, ammonia and inorganic phosphate excretion, and nutrient regeneration of the common jellyfish Aurelia aurita was investigated from 1982 to 1984 in the Kiel Bight, western Baltic Sea. During summer 1982, medusae abundance ranged between 14 and 23 individuals 100 m^-3, biomass was estimated at about 5 g C 100 m^-3 and the mean final diameter of individuals was 22 cm. Abundance, based on numbers, in 1983 and 1984 was an order of magnitude lower; biomass was less than 2 g C 100 m^-3 and jellyfish grew to 30 cm. During the summers of 1983 and 1984, A. aurita biomass constituted roughly 40% of that of the total zooplankton>200 m. In 1982, for which zooplankton data were lacking, it was assumed that medusae biomass was greater than that of all other zooplankton groups. Total ammonia excretion ranged between 6.5 and 36 mol h^-1 individual^-1, whereas inorganic phosphate release was 1.4 to 5.7 mol h^-1 individual^-1. Allometric equations were calculated and exponents of 0.93 for NH₄–N release and 0.87 for PO₄–P excretion were determined. Nitrogen and phosphorus turnover rates were 5.4 and 14.6% d^-1, respectively. In 1982, the medusae population released 1 100 mol NH₄–N m^-2 d^-1, about 11% of the nitrogen requirements of the phytoplankton. The inorganic phosphate excretion (150 mol m^-2 d^-1) sustained 23% of the nutrient demands of the primary producers. In the other two years the nutrient cycling of the medusae was much less important, and satisfied only 3 to 6% of the nutrient demands. It is suggested that in some years A. aurita is the second most important source of regenerated nutrients in Kiel Bight, next to sediment.     

Effects of light intensity on nitrate and nitrite uptake and excretion by Chaetoceros curvisetus

Michaelis-Menten uptake kinetics were observed at all light intensities. With constant illumination, the V_max and K₁ in nitrate uptake over the natural light intensity range of 0 to 2000 E were 0.343 g-at NO₃–N(g)^-1 at protein-N h^-1 and 26 E, respectively. Nitrate uptake was inhibited at higher light intensities. The K_s for nitrate uptake did not vary as a function of light intensity remaining relatively constant at 0.62 g-at NO₃–N 1^-1. With intermittent illumination, the V_mzx for light intensity in nitrate uptake over a light intensity range of 0 to 5000 E was 0.341 g-at NO₃–N(g)^-1-at protein-N h^-1. No inhibition of nitrate uptake was observed at higher than natural light intensities. Chaetoceros curvisetus will probably never experience light inhibition of nitrate uptake under natural conditions.     

Metabolism and elemental composition of zooplankton from the Barents Sea during early Arctic summer

Rates of oxygen consumption, ammonia excretion and phosphate excretion were measured on a hydromedusae (Aglantha digitale), pteropods (Limacia helicina, Clione limacina), copepods (Calanus finmarchicus, C. glacialis, C. hyperboreus, Metridia longa), an amphipod (Parathemisto libellula), a euphausiid (Thysanoessa inermis) and a chaetognath (Sagitta elegans), all of which were dominant species in the Barents Sea during early summer 1987. Water and ash contents and elemental composition (C and N) were also analysed on the specimens used in these metabolic experiments. Between species variations were 67.8% to 94.7% of wet weight in water content, 6.4% to 56.5% of dry weight in ash content, 16.7% to 61.0% of dry weight in carbon content, and 4.3% to 11.2% of dry weight in nitrogen content. Oxygen consumption rates ranged from 0.33 to 13.8 l O₂ individual^-1 h^-1, ammonia excretion rates, from 0.0072 to 0.885 gN individual^-1 h^-1 and phosphate excretion rates, from 0.0036 to 0.33 g P individual^-1 h^-1. In general, higher rates were associated with larger species, but considerable differences were also seen between species. The ratios between the rates (O : N, N : P, O : P) exhibited a wide species-specific variation, indicating differences in dominant metabolic substrates. Typical protein oriented metabolism was identified only in S. elegans. From the results of metabolic rate measurements and elemental analyses, daily losses of body carbon and nitrogen were estimated to be 0.50 to 4.15% and 0.084 to 1.87%, respectively, showing faster turnover rates of carbon than that of nitrogen. Comparison of daily loss of body carbon of the Barents Sea zooplankton with that of the Antarctic zooplankton indicated reduced rates of the former (63% on average).