Energy content at metamorphosis and growth rate of the early juvenile barnacle<Emphasis Type="Italic"> Balanus amphitrite</Emphasis>

The energetic cost of metamorphosis in cyprids of the barnacle Balanus amphitrite Darwin was estimated by quantification of lipid, carbohydrate and protein contents. About 38–58% (4–5 mJ individual^–1) of cypris energy reserves were used during metamorphosis. Lipids accounted for 55–65%, proteins for 34–44% and carbohydrates for <2% of the energy used. Juveniles obtained from larvae fed 10⁶ cells ml^–1 of Chaetoceros gracilis were bigger (carapace length: 560–616 µm) and contained more energy (5.56±0.10 mJ juvenile^–1) than their counterparts (carapace length: 420–462 µm; energy content: 2.49±0.20 mJ juvenile^–1) obtained from larvae fed 10⁴ cells ml^–1. At water temperatures of 30°C and 24°C and food concentrations of 10⁴ and 10² cells ml^–1 (3:1 mixture of C. gracilis and Isochrysis galbana) as well as under field conditions (26.9±3.1°C and 2.2±0.8 µg chlorophyll a l^–1), juveniles obtained from larvae fed the high food concentration grew faster than juveniles obtained from larvae fed low food concentration until 5 days post-metamorphosis. Laboratory experiments revealed a combined effect of early juvenile energy content, temperature and food concentration on growth until 5 days post-metamorphosis. After 10 days post-metamorphosis, the influence of the early juvenile energy content on growth became negligible. Overall, our results indicate that the energy content at metamorphosis is of critical importance for initial growth of juvenile barnacles and emphasize the dependency of the physiological performance of early juvenile barnacles on the larval exposure to food.Communicated by O. Kinne, Oldendorf/LuheAn erratum to this article can be found at     

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     

Otolith microstructure and daily increment validation of marbled sole (<Emphasis Type="Italic">Pseudopleuronectes yokohamae</Emphasis>)

We examined the daily deposition of otolith increments of marbled sole (Pseudopleuronectes yokohamae) larvae and juveniles by rearing experiments, and estimated the growth pattern of wild larvae and juveniles in Hakodate Bay (Hokkaido Island, Japan). At 16°C, prominent checks (inner checks; ca. 19.8 µm in diameter) were observed on the centers of sagittae and lapilli extracted from 5-day-old larvae. On both otoliths, distinctive and regular increments were observed outside of the inner checks, and the slopes of regression lines between age and the number of increments (n_i) (for sagittae: n_i=0.98×Day–5.90; for lapillus: n_i=0.96×Day–5.70) did not significantly differ from 1. Inner check formations were delayed at lower temperature, and the inner checks formed 13 days after hatching at 8°C. Over 80% of larvae, just after their yolk-sac has been absorbed completely (stage C), had inner checks on both their otoliths. On the lapilli, other checks (outer check) formed at the beginning of eye migration (stage G). To validate the daily deposition of increments during the juvenile stage, wild captured P. yokohamae juveniles were immersed in alizarin complexone (ALC)-seawater solutions and reared in cages set in their natural habitat. After 6 days, the mean number of rings deposited after the ALC mark was 5.7. The age–body length relationship of wild P. yokohamae larvae and juveniles caught in Hakodate Bay was divided into three phases. In the larval period, the relationship was represented by a quadratic equation (notochord length=–0.010×Age²+0.682×Age–2.480, r²=0.82, P<0.001), and the estimated instantaneous growth was 0.38 mm day^–1 at 15 days, 0 mm day^–1 at 34 days and –0.12 mm day^–1 at 40 days. The age–body length relationship in the early juvenile stage (<50 days) and the late juvenile stage (>50 days) were represented by linear equations (standard length=0.055×Age+5.722 and standard length=0.345×Age–9.908, respectively). These results showed that the growth rates in the late larval periods and the early juvenile stage were lower than those in the early larval stage and late juvenile stage; during the slow growth period, energy appears to be directed towards metamorphosis rather than body growth. This study provided the information needed to use otolith microstructure analysis for wild marbled sole larvae and juveniles.Communicated by T. Ikeda, Hakodate     

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     

Respiration rate and cost of swimming for Antarctic krill,<Emphasis Type="Italic"> Euphausia superba</Emphasis>, in large groups in the laboratory

Constructing realistic energy budgets for Antarctic krill, Euphausia superba, is hampered by the lack of data on the metabolic costs associated with swimming. In this study respiration rates and pleopod beating rates were measured at six current speeds. Pleopod beating rates increased linearly with current speed, reaching a maximum of 6 beats s^–1 at 17 cm s^–1. There was a concomitant linear increase in respiration rate, from 1.8 mg O₂ g_D^–1 h^–1 at 3 cm s^–1 to 8.0 mg O₂ g_D^–1 h^–1 at 17 cm s^–1. The size of the group tested (50, 100 and 300 krill) did not have a significant effect on pleopod beating rates or oxygen consumption (ANCOVA, F=0.264; P>0.05). The cost of transport reached a maximum of 75 J g^–1 km^–1 at 5 cm s^–1, and then decreased with increasing current speed to 29 J g^–1 km^–1. When considered in light of energy budgets for E. superba, these data indicate that the cost of swimming could account for up to 73% of total daily metabolic expenditure during early summer.Communicated by G.F. Humphrey, Sydney     

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.     

Growth and herbivory by heterotrophic dinoflagellates in the Southern Ocean,studied by microcosm experiments

Growth and herbivory of heterotrophic dinoflagellates (Gymnodinium sp.) from the Weddell Sea and the Weddell/Scotia Confluence were studied in 1988 in 100-liter microcosms. The microcosms were screened through 200-µm or 20-µm mesh nets and incubated for 12 d at 1 °C under artificial light. Mean cell volume of dinoflagellates was 1 000 to 1 500µm³, and that of their phytoplankton prey 360 to 430µm³. Dinoflagellate growth rate followed a Holling type II functional response, with a maximum growth rate of 0.3 d^–1 and half-saturation food concentrations of 1.0µg chlorophylla l^–1, 50µg C l^–1, or 1 500 cells ml^–1. Carbon budgets based on¹⁴CO₂ assimilation and biomasses of phytoplankton and heterotrophic dinoflagellates suggested a balance between phytoplankton grazing loss and dinoflagellate consumption, assuming a dinoflagellate carbon conversion efficiency of 40%. Applying this to the functional response yielded estimates of maximum ingestion rate (0.8µg Cµg^–1 C d^–1, or 6 pg C dinoflagellate^–1 h^–1) and maximum clearance (0.8 to 1.2 × 10⁵ body volumes h^–1, or 80 to 120 nl ind.^–1 h^–1). The microcosm experiments suggested that heterotrophic dinoflagellates may contribute significantly to maintenance of low phytoplankton biomass in the Southern Ocean.     

Food limitation stimulates metamorphosis of competent larvae and alters postmetamorphic growth rate in the marine prosobranch gastropodCrepidula fornicata

The effects of food limitation on growth rates and survival of marine invertebrate larvae have been studied for many years. Far less is known about how food limitation during the larval stage influences length of larval life or postmetamorphic performance. This paper documents the effects of food limitation during larval development (1) on how long the larvae ofCrepidula fornicata (L.) can delay metamorphosis in the laboratory after they have become competent to metamorphose and (2) on postmetamorphic growth rate. To assess the magnitude of nutritional stress imposed by different food concentrations, we measured growth rates (as changes in shell length and ash-free dry weight) for larvae reared in either 0.45-m filtered seawater or at phytoplankton concentrations (Isoehrysis galbana, clone T-ISO) of 1 × l0³, 1 × 10⁴, or 1.8 × 10⁵ cells ml^–1. Larvae increased both shell length and biomass at 1 × 10⁴ cells ml^–1, although significantly more slowly than at the highest food concentration. Larvae did not significantly increase (p > 0.10) mean shell length in filtered seawater or at a phytoplankton concentration of only 1 × 10³ cells ml^–1, and in fact lost weight under these conditions. To assess the influence of food limitation on the ability of competent individuals to postpone metamorphosis, larvae were first reared to metamorphic competence on a high food concentration ofI. galbana (1.8 × 10⁵ cells ml^–1). When at least 80% of subsampled larvae were competent to metamorphose, as assessed by the numbers of indlviduals metamorphosing in response to elevated K⁺ concentration in seawater, remaining larvae were transferred either to 0.45-m filtered seawater or to suspensions of reduced phytoplankton concentration (1 × 10³, 1 × 10⁴, or 5 × 10⁴ cells ml^–1), or were maintained at 1.8 × 10⁵ cells ml^–1. All larvae were monitored daily for metamorphosis. Individuals that metamorphosed in each food treatment were transferred to high ration conditions (1.8 × 10⁵ tells ml^–1) for four additional days to monitor postmetamorphic growth. Competent larvae responded to all food-limiting conditions by metamorphosing precociously, typically 1 wk or more before larvae metamorphosed when maintained at the highest food ration. Surprisingly, juveniles reared at full ration grew more slowly if they had spent 2 or 3 d under food-limiting conditions as competent larvae. The data show that a rapid decline in phytoplankton concentration during the larval development ofC. fornicata stimulates metamorphosis, foreshortening the larval dispersal period, and may also reduce the ability of postmetamorphic individuals to grow rapidly even when food concentrations increase.     

Automatic potentiometric determination of dissolved oxygen

Commercially available automatic titration systems were tested in 1988 for potentiometric titration of liberated iodine by the Winkler method of oxygen determination. The potentiometric equivalence point was also compared to the manual starch end point. Finally the automatic method was used in order to estimate belowhalocline respiration in the Kattegat, Sweden. Standard deviations of 0.007 ml O₂ l^–1 or 0.1 to 0.3% coefficients of variation (% standard deviation of the mean) were achieved when titrating 25 ml from replicate 60-ml oxygen bottles using the automatic method, or 50 ml manually. The precision for replicate titrations of 50-ml aliquots of 0.001N KIO₃ was <0.05% (0.002 ml 0.01N Na₂S₂O₃) for the automatic method. Titration time for 25-ml aliquots was 2 to 4 min, somewhat longer than for manual titrations (1 to 1.5 min). However, during titration the operator is free to perform other tasks. It is not possible to use automatic sample changers, due to rapid iodine volatilization. The equipment can be handled by relatively unskilled analysts and is suitable for use on board research vessels or in field stations [weight for a Mettler^TM titrator (Mettler Instrumente AG, Greifensee, Switzerland) <10 kg, volume <0.1 m³]. Below-halocline oxygen consumption in the SE Kattegat ranged from 0 to 6 ml O₂ m^–3 h^–1 (mean values for September and October 1988=1.69 and 0.66 ml O₂ m^–3 h^–1, respectively, with 95% confidence limits of ca. ±0.6 ml O₂ m^–3 h^–1).     

Ecological and biological features of a mesopelagic ostracod,Conchoecia pseudodiscophora,in the Japan Sea

The abundance and vertical distribution pattern of a halocyprid ostracod,Conchoecia pseudodiscophora, were investigated in the Japan Sea in 1985, 1987 and 1989. Vertical sampling from 500 m depth to the surface in the water around Yamato Rise revealed that this ostracod was second in dominance by number and third to fourth by biomass of the total zooplankton collected with a 0.35 mm mesh Norpac net. Horizontal net tows in Toyama Bay indicated that the major population ofC. pseudodiscophora was distributed below 250 to 300 m depth. No diel migration pattern was evident. Its contribution to total zooplankton there was 5 to 10% or more in terms of biomass. A total of five subadult instars (II to VI) and adult males and females were identified from instar analysis based on sizes and morphological characteristics of specimens collected with 0.10 mm mesh Norpac nets. Data on body length, wet weight and dry weight of each instar are presented. Carbon content of 35 to 48% of dry weight, and nitrogen content of 5.3 to 7.3% of dry weight, were recorded on fresh, freeze-dried specimens of selected instars (subadult Instars IV to VI, adult females). Water and ash contents of mixed specimens of these four instars were 76% of wet weight and 25% of dry weight, respectively. Feasibility of laboratory maintenance ofC. pseudodiscophora was tested, and it produced characteristic J shaped faecal pellets. Oxygen consumption rates of subadult instars V and VI, and adult female ranged 0.011 to 0.021µl O₂ ind.^–1 h^–1 at 1 °C, or 2.9 to 6.1µl O₂ (mg body N)^–0.85 h^–1 in terms of Adjusted Metabolic Rate (AMR_{o 2}). There was no appreciable metabolic reduction inC. pseudodiscophora compared to other ostracods, despite their mesopelagic life mode. Subdominance in total zooplankton and nonreduced metabolic activity ofC. pseudodiscophora suggest that this species may be an important link in mesopelagic energy-flow and matter cycling in the Japan Sea.     

Temporal variability in zooplankton prey capture rate of the passive suspension feeder<Emphasis Type="Italic"> Leptogorgia sarmentosa</Emphasis> (Cnidaria: Octocorallia), a case study

There is increasing evidence that suspension feeders play a significant role in plankton–benthos coupling. However, to date, active suspension feeders have been the main focus of research, while passive suspension feeders have received less attention. To increase our understanding of energy fluxes in temperate marine ecosystems, we have examined the temporal variability in zooplankton prey capture of the ubiquitous Mediterranean gorgonian Leptogorgia sarmentosa. Prey capture was assessed on the basis of gut content from colonies collected every 2 weeks over a year. The digestion time of zooplankton prey was examined over the temperature range of the species at the study site. The main prey items captured were small (80–200 µm), low-motile zooplankton (i.e. eggs and invertebrate larvae). The digestion time of zooplankton prey increased when temperature decreased (about 150% from 21°C to 13°C; 15 h at 13°C, 9 h at 17°C, and 6 h at 21°C), a pattern which has not previously been documented in anthozoans. Zooplankton capture rate (prey polyp^–1 h^–1) varied among seasons, with the greatest rates observed in spring (0.16±0.02 prey polyp^–1 h^–1). Ingestion rate in terms of biomass (g C polyp^–1 h^–1) showed a similar trend, but the differences among the seasons were attenuated by seasonal differences in prey size. Therefore, ingestion rate did not significantly vary over the annual cycle and averaged 0.019±0.002 g C polyp^–1 h^–1. At the estimated ingestion rates, the population of L. sarmentosa removed between 2.3 and 16.8 mg C m^–2 day^–1 from the adjacent water column. This observation indicates that predation by macroinvertebrates on seston should be considered in energy transfer processes in littoral areas, since even species with a low abundance may have a detectable impact.Communicated by S.A. Poulet, Roscoff     

Physiological and biochemical aspects of embryonic and larval development of the winter flounder Pseudopleuronectes americanus

Eggs and larvae of the winter flounder Pseudopleuronectes americanus Walbaum were hatched and raised in the laboratory under controlled conditions. Biochemical composition was measured during development and found to be similar to that of other species: 65 to 80 percent protein, 15 to 30 percent fat, and 0 to 5 percent carbohydrate. Ash content was 7 to 10 percent of dry weight. The chorion comprised more than half of the weight of an egg and the data suggested that it was possibly a source of nutrition to the developing embryo. The sequence of utilization appeared to be carbohydrate and then protein to hatching, lipid, mixed lipid and protein, the predominantly protein until feeding began. Carbohydrate was accumulated at first feeding and depleted when growth began. Protein and lipid were deposited in approximately constant proportions. Respiration rates of eggs were low, 0.002–0.015 l O₂ egg^-1 h^-1, but rose gradually from fertilization to hatching. Respiration rates of early larvae were from two to eight times that of eggs (0.033–0.131 l O₂ larva^-1 h^-1). Variation in larval respiration rates indicated a three-fold difference in rate according to level of activity. Eggs excreted ammonia at an increasing rate from fertilization to hatching. Larvae excreted ammonia, primary amines, and other unidentified organic nitrogenous substances. Rates of excretion and proportions of excretory products varied with stage of development. Primary amine excretion was variable and a major component in early stages. Ammonia-N excreted was two to 20 times primary amine N excreted. Unidentified substances were the predominant form of N excretion during early feeding. Ammonia accounted for most of the N excreted in older larvae. Early specific growth rates were 2.1 and 5.5%. Net caloric conversion and net and gross nitrogen efficiencies were low in first feeding larvae compared to adult fishes (32.2, 27.7, and 10.7% respectively).Contribution no. 5071 from the Woods Hole Oceanographic Institution     

Seasonal and depth related variation in the photosynthesis–irradiance response of<Emphasis Type="Italic"> Ecklonia radiata</Emphasis> (Phaeophyta,Laminariales) at West Island,South Australia

The photosynthesis–irradiance response of Ecklonia radiata (C. Agardh) J. Agardh, a common kelp in the temperate southern hemisphere, was investigated in situ throughout the year and across a depth profile at West Island, South Australia. Temperature and irradiance environment altered throughout the year, varying at 3 m between 14–20°C and 279–705 mol photons m^–2 s^–1. Photosynthetic capacity (P_m) varied throughout the year between 177–278 mol O₂ g^–1 dry wt h^–1 at 3 m and 133–348 mol O₂ g^–1 dry wt h^–1 at 10 m. The irradiance required for sub-saturation of photosynthesis (E_k) varied between 97–152 and 81–142 mol photons m^–2 s^–1 for 3 m and 10 m respectively, and the respiration rate varied between 15–36 and 13–20 mol O₂ g^–1 dry wt h^–1 for 3 m and 10 m. A clear seasonal change in photokinetic parameters was detected and provided strong evidence for a seasonal acclimation response. During winter an increase in the efficiency of light utilisation at low irradiance () was accompanied by a decrease in both E_k and that required for photosynthetic compensation. P_m also increased during the winter and autumn months and respiratory requirements decreased. These changes enable E. radiata to display an optimal photosynthetic performance throughout the year despite significant changes in the surrounding environment.Communicated by P.W. Sammarco, Chauvin     

The influence of macrofauna on estuarine benthic community metabolism: a microcosm study

Effects of benthic macrofauna (Corophium volutator, Hydrobia sp., Nereis virens) on benthic community metabolism were studied over a 65-d period in microcosms kept in either light/dark cycle (L/D-system) or in continuous darkness (D-system). Sediment and animals were collected in January 1986 in the shallow mesohaline estuary, Norsminde Fjord, Denmark. The primary production in the L/D-system after 10 d acted as a stabilizing agent on the O₂ and CO₂ flux rates, whereas the D-system showed decreasing O₂ and CO₂ flux throughout the period. Mean O₂ uptake over the experimental period ranged from 0.38 to 1.24 mmol m^–2 h^–1 and CO₂ release varied from 0.80 to 1.63 mmol m^–2 h^–1 in both systems. The presence of macrofauna stimulated community respiration rates measured in darknes, 1.4 to 3.0 and 0.9 to 2.0 times for O₂ and CO₂, respectively. In contrast, macrofauna lowered primary production. Gross primary production varied from 1.06 to 2.26 mmol O₂ m^–2 h^–1 and from 1.26 to 2.62 mmol CO₂ m^–2 h^–1. The community respiratory quotient (CRQ, CO₂/O₂) was generally higher in the begining of the experiment (0–20 d, mean 1.89) than in the period from Days 20 to 65 (mean 1.38). The L/D-system exhibited lower CRQ (ca. 1) than the D-system. The community photosynthetic quotient varied for both net and gross primary production from 0.64 to 1.03, mean 0.81. The heterotrophic D-system revealed a sharp decrease in the sediment content of chlorophyll a as compared to the initial content. In the autotrophic L/D-system, a significant increase in chlorophyll a concentration was observed in cores lacking animals and cores with C. volutator (The latter species died during the experiment). Due to grazing and other macrofauna activities other cores of the L/D-system exhibited no significant change in chlorophyll a concentration. Community primary production was linearly correlated to the chlorophyll a content in the 0 to 0.5 cm layer. Fluxes of DIN (NH₄ ⁺+NO₂ ^–+NO₃ ^–) did not reveal significant temporal changes during the experiment. Highest rates were found for the cores containing animals, mainly because of an increased NH₄ ⁺ flux. The release of DIN decreased significantly due to uptake by benthic microalgae in the L/D-system. No effects of the added macrofauna were found on particulate organic carbon (POC), particulate organic nitrogen (PON), total carbon dioxide (TCO₂) and NH₄ ⁺ in the sediment. The ratio between POC and PON was nearly constant (9.69) in all sediment dephts. The relationship between TCO₂ and NH₄ ⁺ was more complex, with ratios below 2 cm depth similar to those for POC/PON, but with low ratios (3.46) at the sediment surface.     

Orthophosphate uptake by phytoplankton and bacterioplankton from the Los Angeles Harbor and Southern California coastal waters

Orthophosphate (P) uptake on a seasonal basis in surface waters and in vertical profiles was directly proportional to the standing stocks of phytoplankton and bacterioplankton in the outer Los Angeles Harbor and in southern California coastal waters during 1978–1979. A phytoplankton-enriched size fraction (PEF) which was retained on a 1 m pore-size filter contained 83% of the total chlorophyll a but only 18% of the total bacteria. A bacterioplankton-enriched size fraction (BEF) which passed the 1 m filter but was retained on a 0.2 m filter contained 82% of the total bacteria but only 17% of the total chlorophyll a. PEF and BEF accounted for 91 and 9% of the microbial carbon, respectively. The differential uptake of 10 radiolabeled substrates more fully characterized PEF and BEF. ³³P uptake occurred in both PEF and BEF, accounting for 47 and 53%, respectively, of the total uptake. ³³P uptake by both size fractions was inhibited by low concentrations of 2,4-dinitrophenol (DNP), N-ethylmaleimide (NEM) and carbonyl cyanide, m-chlorophenylhydrozone (CCCP). Darkness and low levels of 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU) selectively inhibited ³³P uptake by PEF; valinomycin selectively inhibited ³³P uptake by BEF. An experiment measuring ³³P uptake velocity versus P concentration produced sigmoidal saturation kinetics at high levels of exogenous P. Kinetic parameter analyses according to the Hill equation gave a V _max of 7.12 nmol l^–1 h^–1 and aK _t of 0.41 nmol l^–1 for PEF, and a V _max of 5.17 nmol l^–1 h^–1 and aK _t of 112 nmol l^–1 for BEF. Consideration of relative surface areas of phytoplankton and bacterioplankton, their ³³P uptake rates in light and dark, and estimates of the population turnover times emphasizes the potential importance of bacterioplankton in community phosphorus metabolism.     

Respiration,photosynthesis and carbon metabolism in planktonic ciliates

Release of¹⁴C-labelled carbon dioxide from uniformly labelled cells was used to measure respiration by individual ciliates in 2-h incubations in 1989 and 1990. In a strictly heterotrophic ciliate,Strobilidium spiralis (Leegaard, 1915), release of labelled carbon dioxide was equivalent to ca. 2.8% of cell C h^–1 at 20°C, and there was no difference between rates in the dark and light. In the chloroplast-retaining ciliatesLaboea strobila Lohmann, 1908,Strombidium conicum (Lohmann, 1908) Wulff, 1919 andStrombidium capitatum (Leegaard, 1915) Kahl, 1932, release of labelled carbon dioxide was less in the light than in the dark in experiments done at 15°C. InL. strobila release of radiolabel as carbon dioxide was equivalent to ca. 2.4% of cell C h^–1 in the dark but ca. 1% at 50µE m^–2 s^–1, an irradiance limiting to photosynthesis. InS. conicum release of radiolabel as carbon dioxide was equivalent to ca. 4.4% of cell C h^–1 in the dark, but at an irradiance saturating to photosynthesis (250 to 300µE m^–2 s^–1) there was no detectable release of labelled carbon dioxide. InS. capitatum release of radiolabel as carbon dioxide was equivalent to ca. 4.3% of cell C h^–1 in the dark but at an irradiance saturating to photosynthesis was ca. 2.4% of cell C h^–1. These data, combined with data from photosynthetic uptake experiments, indicate that¹⁴C uptake underestimates the total benefit of photosynthesis by 50% or more in chloroplastretaining ciliates.Contribution no. 7510 from the Woods Hole Oceanographic Institution     

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     

Reproductive cycle,larval development,juvenile growth and population dynamics of Patiriella pseudoexigua (Echinodermata: Asteroidea) in Taiwan

A field study was conducted at Wanlitung, southern Taiwan, in 1986–1089, to determine the reproductive cycle, development mode, growth rate and population dynamics of the small seastar Patiriella pseudoexigua (Dartnall), which occurs in highly stressful and disturbed intertidal pools in this area. An inverse relationship between gonad index and pyloric-caccum index was only recorded immediately prior to spawning. A short, well-synchronized seasonal spawning occurs in October. When reared at 25 °C, lecithotrophic larvae develop directly, lack a bipinnaria stage, and metamorphose completely on the seventh day after fertilization. The growth curves of field juveniles are linear, those of laboratory-reared juveniles are sigmoid. Juveniles appear in tide pools in spring-early summer of each year. Adults spawn mainly in late fall, enabling spawning to occur in time for the larvae to benefit from the environmentally favorable winter season. Populations in high-tidal pools decrease in later summer, but remain more stable in lowtidal pools and lagoons.     

An energy budget for the free-swimming and metamorphosing larvae of Balanus balanoides (Crustacea: Cirripedia)

Analysis of biochemical components and measurements of oxygen consumption rates of cypris larvae of Balanus balanoides (L.) maintained in the laboratory at 10°C for up to 5 weeks after capture shows that lipid is the primary energy reserve, although later protein is utilised. Initially, the cyprids swim freely with an oxygen consumption rate of ca. 37×10^-3 l O₂ h^-1 cyprid^-1, but within a few days the rate falls to ca. 21×10^-3 l O₂ h^-1 cyprid^-1 when they cease swimming and explore the substratum. The cost of metamorphosis was calculated both from the loss of biochemical components and oxygen consumption rates during metamorphosis; the values were 2.8×10^-2 and 3.2×10^-2 cal cyprid^-1, respectively. A budget was collated from the data on respiration and biochemical composition, whereby the energy per cyprid was partitioned into that required for essential structural components (6.8×10^-2 cal), that needed for metamorphosis (3.0×10^-2 cal) and an excess available for swimming and exploring, which in the batches studied was about 5.0×10^-2 cal. This excess is mainly derived from the utilisation of lipid reserves and is used up usually 2 1/2 to 4 weeks after capture. During these measurements, samples of cyprids were taken at weekly intervals to test the rate of settlement and success of metamorphosis. The results showed that they lose their competence to metamorphose successfully approximately at the same time (3 to 4 weeks) that the energy supply for swimming and exploration is used up.     

Interactive accumulation of mercury and selenium in the sea starAsterias rubens

Interactions between mercury and selenium accumulation and subcellular binding inAsterias rubens (L.), collected in 1987 from Lille Bælt at Middelfart, Funen, Denmark, were investigated in laboratory experiments. Sea stars exposed to 10µg Hg l^–1 for 30 d accumulated mercury in body wall, tube feet and stomach linearly with time at 1.2, 1.2 and 0.5µg Hg g^–1 dry wt d^–1, respectively. Mercury was accumulated in pyloric caeca and coelomic fluid initially at 1.4µg Hg g^–1 dry wt d^–1 and 9.4 ng Hg ml^–1 d^–1, respectively; after 10 d uptake rates decreased. Sea stars exposed to 75µg Se-SeO ₃ ^{- -} l^–1 accumulated selenium linearly with time over 30 d in the stomach, pyloric caeca, tube feet and body wall at 2.0, 1.2, 1.2 and 0.6µg Se g^–1 dry wt d^–1. Sea stars exposed to 75µg Se-SeO ₄ ^{- -} l^–1 maintained selenium levels in the coelomic fluid at 75µg Se l^–1 over 30 d. Exposure to selenate did not alter the selenium concentrations in the tissues. Sea stars exposed concurrently to 75µg Se-SeO ₃ ^{- -} and 10µg Hg l^–1 accumulated more mercury and selenium in tube feet and body wall than did sea stars exposed to the two elements alone. In pyloric caeca and stomach concurrent exposure reduced accumulation of both elements. Mercury was bound predominantly in the insoluble fraction of the tissues, and soluble mercury was bound in proteins of high (> 70 kilodaltons) or very low (< 6000 daltons) molecular weight. Ca. half of the selenium recovered was bound in the insoluble fraction, and soluble selenium was bound in proteins of high (> 70 kilodaltons) or very low (< 6000 daltons) molecular weight. Interaction between the two elements was exerted predominantly in the insoluble fraction of the tissues.