Growth and grazing responses of tintinnid ciliates feeding on the toxic dinoflagellate Heterocapsacircularisquama

Growth and feeding rates of two tintinnid species, Favellaazorica and Favellataraikaensis, were determined under various concentrations of the dinoflagellate Heterocapsacircularisquama which has been reported as highly toxic to shellfish. Mean growth rates of F. azorica and F. taraikaensis on a diet of H.circularisquama (ca. 10² cells ml⁻¹) were 2.15 and 1.97 doublings d⁻¹, respectively. These values are similar to those on a diet of Heterocapsatriquetra which is suitable food for various zooplankton. However, growth rates of both tintinnid species decrease with increasing concentrations of >10³ cells ml⁻¹ of H. circularisquama. In particular, H. circularisquama under conditions of >10³ cells ml⁻¹ caused mortality in F.taraikaensis, probably due to toxins. Clearance and ingestion rates of F. azorica on H. circularisquama were 4.1 to 27.5 μl ind⁻¹ h⁻¹ and 1.5 to 28.7 cells ind⁻¹ h⁻¹, respectively, at concentrations of <10⁴ cells ml⁻¹ and those of F. taraikaensis were 0.9 to 22.1 μl ind⁻¹ h⁻¹ and 0.1 to 13.0 cells ind⁻¹ h⁻¹, respectively, at concentrations of <10³ cells ml⁻¹. Both clearance and ingestion rates on H.circularisquama were higher for replicates fed on H.triquetra. Daily grazing impact of the two species of Favella on the initial stage of a bloom of H.circularisquama were estimated to reach 6 to 50% of H. circularisquama at a concentration of 540 cells ml⁻¹, indicating that grazing by tintinnids such as Favella spp. may significantly regulate the initial stages of blooms of H. circularisquama. Received: 3 January 1997 / Accepted: 27 January 1997     

Growth and grazing rates of the tintinnid ciliate<Emphasis Type="Italic"> Favella taraikaensis</Emphasis> on the toxic dinoflagellate<Emphasis Type="Italic"> Alexandrium tamarense</Emphasis>

Growth and feeding activities of the tintinnid ciliate Favella taraikaensis fed the toxic dinoflagellate Alexandrium tamarense were examined in laboratory experiments. Both growth and ingestion rates of F. taraikaensis as a function of the A. tamarense concentration were fitted to a rectangular hyperbolic equation. The maximum growth and ingestion rates of F. taraikaensis were 1.0 day^–1 and 2.8 cells ind. h^–1 (carbon specific ingestion rates: 3.5 day^–1), respectively, which are both included in the range of previous data reported for Favella spp. feeding on other algae. The gross growth efficiency (GGE) of F. taraikaensis ranged from 0.26 to 0.49 (mean value 0.40) at the concentration of 10–800 cells ml^–1, which is within the range of previous data on Favella spp. Also, the growth and ingestion rates and GGE of F. taraikaensis on A. tamarense were not significantly different from the values on another non-toxic dinoflagellate (Heterocapsa triquetra) at two different prey concentrations. This indicates that the toxicity of A. tamarense probably did not influence the feeding and growth activities of F. taraikaensis at concentrations of less than ca. 800 cells ml^–1. To evaluate the grazing by F. taraikaensis on A. tamarense blooms in the field, the population dynamics of A. tamarense were simulated based on the growth and ingestion parameters of F. taraikaensis. As a result, the grazing impact by F. taraikaensis was considered to potentially regulate the development of A. tamarense blooms. If the toxicity of A. tamarense does not influence the growth and feeding activities of F. taraikaensis, the occurrence of such grazer plankton are considered to be important for predicting the course of A. tamarense bloom dynamics under natural conditions.Communicated by T. Ikeda, Hakodate     

Grazing of Acartia hudsonica (A. clausi) on Skeletonema costatum in Narragansett Bay (USA): Influence of food concentration and temperature

Grazing experiments were performed with temperatureacclimated Acartia hudsonica fed the diatom Skeletonema costatum in concentrations ranging from 50 to 3×10⁴ cell ml^-1 at 5°, 10° and 15°C. The ingestion data were best fit by an Ivlev equation. Feeding threshold values of 39 and 59 cells ml^-1 were not significantly different from zero; however, filtration rates were depressed at low food concentrations. Maximum filtration rates increased exponentially with temperature, reaching a maximum with copepods collected at 14°–15°C, and then declining. Both the increase in ingestion rate with increasing food concentration and the maximum ingestion rate were significantly greater as experimental temperature was increased. Maximum ingestion rates were reached at concentrations greater than 6×10³ cells ml^-1. Percent of body carbon ingested per day at 5 g C L^-1 increased from 1.5% at 5°C to 6.7% at 15°C. At 500 g C L^-1, the ingestion increased from 84% (5°C) to 660% (15°C). Percent of body nitrogen at 0.5 g N L^-1 increased from 0.6% per day at 5°C to 2.5% per day at 15°C. At 50 g N L^-1, the ingestion was 42% body nitrogen at 5°C and 250% at 15°C. The influence of grazing by A. hudsonica on phytoplankton in Narragansett Bay, USA was estimated for 1972–1977. The percent of standing stock removed by grazing rarely exceeded 5% per day except during the late spring when S. costatum growth becomes nutrient limited and higher temperatures favor the rapid population growth of A. hudsonica.     

Ingestion,growth and development of Penaeus indicus larvae as a function of Thalassiosira weissflogii cell concentration

Penaeus indicus larvae have been successfully reared in the laboratory using Thalassiosira weissflogii, Brachionus plicatilis and Artemia salina nauplii as food, with an average survival of 95.8% from nauplius 6 to postlarva 1. The effect of T. weissflogii cell concentration on larval ingestion, development and growth (total length) was investigated. Cell ingestion rates showed a saturation response to concentration. Both maximum ingestion rates and incipient limiting levels (the lowest concentration before ingestion rates were limited) were established for the feeding larval stages. Both were found to increase with progressive increase in larval development. Maximum ingestion rates increased from 0.25×10⁴ cells. larva^-1.h^-1 during protozoea 1 to reach a peak of 1.2×10⁴ cells. larva^-1.h^-1 during mysis 3 and then declined to 0.6×10⁴ cells. larva^-1.h^-1 at postlarva 1. Incipient limiting levels (ILLs) increased from approximately 0.6×10⁴ cells.ml^-1 during protozoea 2, to 0.65×10⁴ cells.ml^-1 during mysis 1, to 1.3×10⁴ cells. ml^-1 during mysis 3 to 1.6×10⁴ cells.ml^-1 at post-larva 1. Filter feeding efficiency was found to reach a maximum during mysis 1. Filter mechanisms are discussed. Generally, the most advanced larval development per unit time occurred at concentrations at and above the ILLs, while retarded development occurred below these levels. Growth increased asymptotically with cell concentration. Incipient growth limiting levels (IGLLs; the lowest concentration before growth was significantly limited) also increased with larval development and with the exception of mysis 3 they coincided with the ILLs. IGLLs increased from 0.55×10⁴ cells.ml^-1 during protozoea 2, to 0.66×10⁴ cells.ml^-1 during mysis 1, to 0.99×10⁴ cells.ml^-1 during mysis 3, to 1.62×10⁴ cells.ml^-1 at postlarva 1. Below the ILLs where ingestion was limited, animals were significantly smaller, with larval development and growth positively correlated to ingestion rates. When culturing penaeoid larvae, ambient cell concentrations should be kept above these known limiting levels to yield consistently good larval survival and growth.     

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     

<Emphasis Type="Italic">Cochlodinium polykrikoides</Emphasis> blooms and clonal isolates from the northwest Atlantic coast cause rapid mortality in larvae of multiple bivalve species

Globally, many commercial bivalve populations have declined in recent decades. In addition to overharvesting and habitat loss, the increasing frequency and intensity of harmful algal blooms (HABs) are likely to contribute to bivalve losses, particularly in cases where blooms negatively impact larval stages. This paper reports on the lethal effects of clonal cultures and blooms of Cochlodinium polykrikoides from the US Atlantic coast on the larvae of three species of commercially and ecologically valuable bivalves: the Eastern oyster (Crassostrea virginica), the bay scallop (Argopecten irradians), and the Northern quahog (hard clam; Mercenaria mercenaria). Both cultures and blooms of C. polykrikoides were highly toxic to all three species of bivalve larvae causing 80–100% mortality during 24- to 72-h exposures at concentrations of 1–2 × 10³ cells ml⁻¹. Toxicity was dependent on cell densities, growth stage of C. polykrikoides (i.e. cultures in exponential stage growth were more toxic than later stages), exposure time of larvae to cells (i.e. longer exposure caused higher mortality), the age of larvae (i.e. younger larvae were more sensitive), and the relative abundance of C. polykrikoides (i.e. the presence of other microalgae decreased toxicity). Free radical-scavenging enzymes (peroxidase and catalase) and the removal of C. polykrikoides cells (i.e. culture filtrate) significantly increased larval survival suggesting toxicity is maximized by contact with live cells and may involve labile toxins bound by these compounds including e.g. reactive oxygen species. The toxicity of C. polykrikoides to bivalve larvae was generally more severe than other HAB species (e.g. Karenia brevis, Karlodinium veneficum, Alexandrium tamarense, Prorocentrum minimum). Since the bivalves in this study spawn in the months when C. polykrikoides blooms on the east coast of North America, these results suggest that these blooms may have detrimental effects on efforts to restore these already diminished populations.     

Effects of the dinoflagellates Karlodinium veneficum and Prorocentrum minimum on early life history stages of the eastern oyster (Crassostrea virginica)

The bloom-forming dinoflagellates Prorocentrum minimum and Karlodinium veneficum can have detrimental effects on some marine life, including shellfish, but little is known about their effects on early life history stages of bivalves. In the Chesapeake Bay region, blooms of these dinoflagellates overlap with the spawning season of the eastern oyster, Crassostrea virginica. In laboratory experiments, we compared the effects of P. minimum and K. veneficum on the survival and development of embryos and larvae of the eastern oyster. At 10⁴ cells ml⁻¹, P. minimum did not have a negative effect on embryos and larvae in 2-day exposures. The yield of D-hinge larvae was equal to or greater than in control treatments. At 2 × 10⁴ cells ml⁻¹ (approximately equal biomass to the P. minimum treatment) K. veneficum caused significant mortality to oyster embryos within 1 day and almost no embryos developed into D-hinge larvae. This effect was not alleviated by the provision of an alternate food source (Isochrysis sp.). Significant mortality was observed when larvae were exposed to K. veneficum at concentrations of 10⁴ cells ml⁻¹ (approximately 5 ng ml⁻¹ of karlotoxin). The K. veneficum cultures used in these experiments were relatively low in toxin content, more toxic strains could be expected to cause mortality at lower cell concentrations. Survival and maturation of embryos and larvae may be reduced when spawns of the eastern oyster coincide with high bloom densities of K. veneficum.     

Influence of food concentration,temperature and salinity on the larval development ofBalanus amphitrite

Influence of food concentration (0.5, 1 and 2 x 10⁵ cell ml^–1 ofSkeletonema costatum), temperature (20 and 30°C) and salinity (15, 25 and 35) on the larval development ofBalanus amphitrite (Cirripedia: Thoracica) was examined. The mortality rate at 20°C was lower than at 30°C in general. Increase in food concentration from 0.5 to 1 x 105 cells ml^–1 improved the survival rate, but this was not evident when food concentration was increased to 2 x 105 cells ml^–1. The results indicate that food availability and temperature jointly determine the energy allocation for metamorphic progress. It was observed that the influence of the tested variables varied with instar. At 20 °C the mean duration of the second instar exceeded 3 d and was much longer than other instar durations. The fourth, fifth and sixth instars and the total naupliar period showed that the effect of different salinities at given food concentrations was negligible at 20°C, while at 30°C there was a marked decrease in duration with increasing salinity.     

Energy budgest,growth and filtration rates in Mytilus edulis at different algal concentrations

Growth of Mytilus edulis L. was measured in aquaria with through-flowing sea water at different levels of constant algal concentrations. The amount of food and oxygen consumed by the mussels were measured over given periods as well as the changes in dry organic weight during the same periods. From these parameters it was possible to make simple energy budgets and to compare the estimated growth with actual growth, and, further, to determine growth efficiences at different food levels. Energy budgets were made for mussels grown at algal concentrations of 0, 1.6×10³, 3.0×10³ and 26.0×10³ Phaeodactylum tricornutum cells x ml^-1. The estimated growth was found to be close to actual growth at algal concentrations above maintenance level and the net growth efficiency was found to be between 18% (3.0×10³ cells x ml^-1) and 61% (26×10³ cells x ml^-1). It has been shown that the filtration rate is independent of algal concentrations between about 1.5×10³ to 30×10³ P. tricornutum cells x ml^-1. Outside this range a decrease in filtration rate was noticed.     

Über den Einfluß der Nahrungskonzentration und anderer Faktoren auf Filtrierleistung und Nahrungsausnutzung der Muscheln Arctica islandica und Modiolus modiolus

Filtration rates and the extent of phagocytosed food particles were determined in the offshore lamellibranchs Artica islandica and Modiolus modiolus in relation to particle concentration, body size and temperature. Pure cultures of the algae Chlamydomonas sp. and Dunaliella sp. were used as food. A new method for determining filtration rates was developed by modifying the classical indirect method. The concentration of the experimental medium (100%) was kept constant to ±1%. Whenever the bivalves removed algae from the medium, additional algae were added and the filtration rate of the bivalves expressed in terms of percentage amount of algae added per unit time. The concentration of the experimental medium was measured continuously by a flow colorimeter. By keeping the concentration constant, filtration rates could be determined even in relation to different definite concentrations and over long periods of time. The amount of phagocytosed food was measured by employing the biuret-method (algae cells ingested minus algae cells in faeces). Filtration rates vary continuously. As a rule, however, during a period of 24 h, two phases of high food consumption alternate with two phases of low food consumption during which the mussels' activities are almost exclusively occupied by food digestion. Filtration rate and amount of phagocytosed algae increase with increasing body size. Specimens of A. islandica with a body length of 33 to 83 mm filter between 0.7 to 71/h (30–280 mg dry weight of algae/24 h) and phagocytose 21 to 122 mg dry weight of algae during a period of 24 h. The extent of food utilization declines from 75 to 43% with increasing body size. In M. modiolus of 40 to 88 mm body length, the corresponding values of filtration rate and amount of phagocytosed algae range between 0.5 and 2.5 l/h (20–100 mg dry weight of algae) and 17 to 90 mg dry weight of algae, respectively; the percentage of food utilization does not vary much and lies near 87%. Filtration rate and amount of phagocytosed algae follow the allometric equation y=a·x ^b. In this equation, y represents the filtration rate (or the amount of phagocytosed algae), a the specific capacity of a mussel of 1 g soft parts (wet weight), x the wet weight of the bivalves' soft parts, and b the specific form of relationship between body size and filtration rate (or the amount of phagocytosed algae). The values obtained for b lie within a range which indicates that the filtration rate (or the amount of phagocytosed algae) is sometimes more or less proportional to body surface area, sometimes to body weight. Temperature coefficients for the filtration rate are in Arctica islandica Q₁₀ (4°–14°C)=2.05 and Q₁₀ (10°–20°C)=1.23, in Modiolus modiolus Q₁₀ (4°–14°C)=2.33 and Q₁₀ (10°–20°C)=1.63. In A. islandica, temperature coefficients for the amount of phagocytosed algae amount to Q₁₀ (4°–14°C)=2.15 and Q₁₀ (10°–20°C)=1.55, in M. modiolus to Q₁₀ (4°–14°C)=2.54 and Q₁₀ (10°–20°C)=1.92. Upon a temperature decrease from 12° to 4°C, filtration rate and amount of phagocytosed algae are reduced to 50%. At the increasing concentrations of 10×10⁶, 20×10⁶ and 40×10⁶ cells of Chlamydomonas/l offered, filtration rates of both mollusc species decrease at the ratios 3:2:1. At 12°C, pseudofaeces production occurs in both species in a suspension of 40×10⁶, at 20°C in 60×10⁶ cells of Chlamydomonas/l. At 12°C and 10–20×10⁶ cells of Chlamydomonas/l, the maximum amount of algae is phagocytosed. At 40×10⁶ cells/l, the amount of phagocytosed cells is reduced by 26% as a consequence of low filtration rates and intensive production of pseudofaeces. At 20°C and 20–50×10⁶ cells of Chlamydomonas/l, the maximum amount of algae is sieved out and phagocytosed; the concentration of 10×10⁶ cells/l is too low and cannot be compensated for by increased activity of the molluscs. With increasing temperatures, the amount of suspended matter, allowing higher rates of filtration and food utilization, shifts toward higher particle concentrations; but at each temperature a threshold exists, above which increase in particle density is not followed by increase in the amount of particles ingested. Based on theoretical considerations and facts known from literature, 7 different levels of food concentration are distinguishable. Experiments with Chlamydomonas sp. and Dunaliella sp. used as food, reveal the combined influence of particle concentration and particle size on filtration rate. Supplementary experiments with Mytilus edulis resulted in filtration rates similar to those obtained for M. modiolus, whereas, experiments with Cardium edule, Mya arenaria, Mya truncata and Venerupis pullastra revealed low filtration rates. These species, inhabiting waters with high seston contents, seem to be adapted to higher food concentrations, and unable to compensate for low concentrations by higher filtration activities. Adaptation to higher food concentrations makes it possible to ingest large amounts of particles even at low filtration rates. Suspension feeding bivalves are subdivided into four groups on the basis of their different food filtration behaviour.     

Effects of algal and larval densities on development and survival of asteroid larvae

Effects of larval and algal culture density and diet composition on development and survival of temperate asteroid larvae were studied in the laboratory at Santa Cruz, California, USA, during summer and fall of 1990. Larvae of Asterina miniata were reared at two densities, 0.5 or 1.0 ml^-1, and fed one or two species of cultured phytoflagellates — Dunaliella tertiolecta alone or mixed with Rhodomonas sp. — at three concentrations of 5x10², 5x10³, and 5x10⁴ total cells ml^-1. Algal concentration strongly influenced larval development; however, larval density also had a marked effect. Development progressed further with increasing algal concentration. Larval growth and differentiation were sometimes uncoupled; i.e., growth measures were directly related to food level, while differentiation indicators were less so. At the lowest food level, growth was negative and differentiation was arrested at early precompetent stages; these larvae never formed juvenile rudiments or brachiolar attachment structures. Development times of larvae given more food ranged from 26 to 50 d and depended directly on food availability. Development time to metamorphosis at the highest food concentration was similar for siblings fed D. tertiolecta alone or mixed with Rhodomonas sp. In contrast, when food level was an order of magnitude lower, larvae fed the algal mixture metamorphosed significantly earlier than larvae fed the unialgal diet. This suggests interactive effects of food quantity and food quality. Survival was little affected by larval or food density, except at the lowest ration. Feeding experiments in well-controlled laboratory conditions are useful to predict and compare the physiological or developmental scope of response of larvae to defined environmental factors; however, results from such studies should not be extrapolated to predict rates and processes of larval development in nature.     

Characteristics of feeding on dinoflagellates by newly hatched larval crabs

The zoeal larvae of brachyuran crabs must feed soon after hatching on a diet that includes large micro- and mesozooplankton in order to satisfy nutritional requirements. However, newly hatched larvae have been shown to ingest a variety of dinoflagellates, perhaps using microbial carbon sources to sustain them until they encounter more favored prey. Ingestion of dinoflagellates by larval crabs has been documented previously under conditions in which the larvae were exposed to algae provided in monoculture or in defined mixtures of cells. We report here on experiments conducted on the hatching stage of five crab species to determine if ingestion of dinoflagellates occurred when they were provided in combination with Artemia sp. nauplii or after a period of feeding on mesozooplankton. Quantitative measurements of chl a in the larval guts provided evidence of ingestion of algal cells. Active ingestion of the dinoflagellate Prorocentrum micans at specified intervals during an extended feeding period was determined on larvae of two crab species using fluorescently labeled cells provided for brief periods at prescribed time intervals. Stage 1 larvae of four of the five crab species ingested dinoflagellates when they were provided in combination with nauplii and larvae of all five species ingested cells after feeding solely on nauplii for 24 h. Ingestion of algal cells was first evident in the larval guts after 6 h of feeding at both low (200 cell ml⁻¹) and high (1,000 cells ml⁻¹) prey densities. Higher prey densities resulted in higher gut chl a. Larvae continuously exposed to dinoflagellates actively ingested cells at every 3 h interval tested over a 36 h period. Results confirm previous studies that larvae will ingest dinoflagellates even when they are encountered in a mixed prey field or when having previously fed. Ingestion of cells may occur on a continual basis over time.     

Vertical distributions and photosynthetic action spectre of two oceanic picophytopianlcters,Prochlorococcus marinus andSynechococcus sp.

Two picophytoplankters,Prochlorococcus marinus andSynechococcus sp., were isolated from the bottom of the euphotic zone (150 m depth) in the western Pacifie Ocean. The concentration ofP. marinus at this depth was more than 10⁴ cells ml^–1 while that ofSynechococcus sp. was less than 10² cells ml^–1. TheP. marinus isolate has a high divinyl-chlorophylla:b ratio similar to that of the Mediterranean strain, while theSynechococcus sp. isolate is of the phycourobilinrich type. The growth rate ofP. marinus was higher thanSynechococcus sp. when both were cultured under weak blue-green to blue-violet light (ca. 2 E m^–2 s^–1). While the chlorophyll-specific absorption spectra showed higher values inSynechococcus sp., the photosynthetic action spectre revealed thatP. marinus was able to use blue-violet light, whereasSynechococcus sp. was able to use blue-green light, more efficiently for photosynthesis. The photosynthetic quantum yield ofP. marinus was higher than that ofSynechococcus sp. at any wavelength between 400 and 700 nm. The calculated in situ photosynthesis rates per Gell volume forP. marinus were estimated to be higher than forSynechococcus sp. at 50 and 150 m depth. These results indicate thatP. marinus photosynthetically surpassesSynechococcus sp. in the blue-light-rieh environment of the oceanic euphotic zone. This may be why the former predominates at depths in temperate to tropical open ocean waters.     

Effects of copper and zinc on two planktonic ciliates

The interactive effects of copper and zinc on two estuarine planktonic ciliates, Favella sp. and Balanion sp., were determined in seawater media in which the free metal ion activities were controlled by nitrilotriacetic acid (NTA) trace metal ion buffer systems. Cupric ion activities of 10^-10 M caused abnormal motility in both ciliates in shortterm (5 h) tests, and cupric ion activities as low as 10^-12.8 M decreased the growth rates of both species in longer-term experiments. In the short-term tests, zinc ion activity by itself did not affect the motility of the ciliates, but there were significant interactions between copper and zinc. In the longer-term experiments, the growth of Favella sp. was optimal at the lowest cupric ion activity (10^-13 M) and the two lowest zinc ion activities (10^-12 and 10^-13 M) and the two lowest zinc ion activities (10^-12 and 10^-11 M), and copper and zinc inhibited growth at activities above these values. By contrast, optimal growth rate of Balanion sp. occurred at the highest zinc ion activity (10^-10 M) and the lowest cupric ion activities (10^-13 to 10^-12 M) and growth rate was reduced at zinc ion activities 10^-11 M. There was an antagonism between copper and zinc which was particularly pronounced in Balanion sp.Contribution No. 5871 from the Woods Hole Oceanographic Institution     

Blooms of the dinoflagellate Gyrodinium aureolum in a Long Island estuary:

A dense dinoflagellate bloom of Gyrodinium aureolum Hulburt in a shallow temperate zone estuary was monitored during the summers of 1982 and 1983. The bloom was typically extremely localized, its densest part exceeding 1000g chlorophyll a liter^-1 (2x10⁴ cells ml^-1). The bloom began at temperatures between 24.5° and 27°C, existed at as high as 30°C and terminated when water temperature dropped to between 19° and 22°C. The highest specific growth rate measured was 0.90d^-1 (1.3 divisions d^-1) and near the termination of the bloom decreased to 0.28d^-1 (0.4 divisions d^-1). A diel vertical migration of the bloom was observed. A box model analysis, based on division rates, vertical migration and water circulation patterns, indicated that the bloom must move downward at the estuary mouth to maintain itself in the estuary, either by means of a convergence system or by downward swimming. High growth rate, low grazing pressure, and a stratified water column are proposed to stimulate bloom formation. Decreasing growth rate appeared to reduce the intensity of the bloom and finally allowed its disappearance by estuarine flushing and mixing.Communicated by J. M. Shick, Orono     

Zooplankton grazing on the coccolithophore Emiliania huxleyi and its role in inorganic carbon flux

Grazing and faecal pellet production by the copepods Calanus helgolandicus and Pseudocalanus elongatus, feeding on the coccolithophore Emiliania huxleyi, were measured under defined laboratory conditions, together with the chemical characteristics and sinking rates of the faecal pellets produced. Ingestion rates of both copepods were equivalent at comparable cell concentrations, the relationship between ingestion rate (I, cells copepod^-1 h^-1) and food concentration (C, cells ml^-1), being I=0.558C for both species. P. elongatus produced a larger number of smaller faecal pellets than C. helgolandicus, but egested a larger volume of material per individual. Only between 27 and 50% of the ingested coccolith calcite was egested in the faecal pellets, and it is possible that acid digestion in the copepod gut is responsible for these considerable losses. Average sinking rates of faecal pellets containing E. huxleyi coccoliths, produced by both species, were >100 m d^-1. The implications of the quantitative laboratory estimates for the vertical flux of inorganic carbon are considered using recently studied shelf-break and oceanic E. huxleyi blooms in the N. E. Atlantic as examples.     

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.     

Two different proteases produced by a deep-sea psychrotrophic bacterial strain,<Emphasis Type="Italic"> Pseudoaltermonas</Emphasis> sp. SM9913

A psychrotrophic bacterial strain, Pseudoaltermonas sp. SM9913, was isolated from deep-sea sediment collected at 1,855 m depth. Two proteases produced by Pseudoaltermonas sp. SM9913 were purified, MPC-01 and MCP-02. MCP-01 is a serine protease with a molecular weight of 60.7 kDa. It is cold-adapted with an optimum temperature of 30–35°C. Its K_m and E_a for the hydrolysis of casein were 0.18% and 39.1 kJ mol^–1, respectively. It had low thermostability, and its activity was reduced by 73% after incubation at 40°C for 10 min. MCP-02 is a mesophilic metalloprotease with a molecular weight of 36 kDa. Its optimum temperature for the hydrolysis of casein was 50–55°C. The K_m and E_a of MCP-02 for the hydrolysis of casein were 0.36% and 59.3 kJ mol^–1, respectively. MCP-02 had high thermostability, and its activity was reduced by only 30.5% after incubation at 60°C for 10 min. At low temperatures, Pseudoaltermonas sp. SM9913 mainly produced the psychrophilic protease MCP-01.Communicated by O. Kinne, Oldendorf/Luhe     

Ecological growth strategies in the seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae): Soluble nitrogen and reserve carbohydrates

The seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae) were grown in an outdoor continuous-flow system at both ambient incident light (I₀) and 0.13 I₀. During the winter, both species accumulated substantial soluble nitrogen reserves (up to 1020 g-at N·g dry wt^-1 in G. foliifera and 630 g-at N·g dry wt^-1 in Ulva sp.). The rate at which these N reserves were depleted was proportional to the growth rate. Seaweeds grown at 0.13 I₀ had lower growth rates and higher levels of soluble tissue N than plants grown at I₀. During the spring-summer growing season, peaks in tissue N followed nutrient peaks in the ambient seawater. Ulva sp. had higher nutrient uptake and growth rates than G. foliifera and showed greater fluctuations in soluble tissue N. This may characterize opportunistic seaweed species with high biomass turnover rates. At I₀, the levels of starch (up to 340 mg·g dry wt^-1 in G. foliifera and 170 mg·g dry wt^-1 in Ulva sp.) were highest during the spring and summer. During this period, fluctuations in starch content were inversely related to growth rate and soluble tissue N. Seaweeds grown at 0.13 I₀ did not accumulate starch. Neither species was found to overwinter with starch reserves.     

Effects of suspended sediments on egg production of the calanoid copepod Acartia tonsa

The estuarine copepod Acartia tonsa was collected on several occasions between 4 April and 14 August 1985 from Terrebonne Bay, Louisiana (29°08N; 90°36W) and the effects in its diet of suspended sediments, collected from the same area, were measured at five different concentrations of sediment (100 to 1 000 ppm) and six phytoplankton concentrations (500 to 13 000 cells ml^-1 Thalassiosira weissflogii). Egg production rate was used as an index of diet quality. At low phytoplankton concentrations (500 cells ml^-1), and at intermediate phytoplankton concentrations (2 000 cells ml^-1) for previously starved copepods, egg production was reduced by up to 40% at a sediment concentration of 250 ppm and further reduced at higher sediment concentrations. At higher food concentrations (4 000 to 13 000 cells ml^-1), suspended sediment had no effect on egg production rates at sediment concentrations up to 500 ppm. Rates were reduced only at the highest sediment concentration of 1 000 ppm. Under most natural conditions, suspended sediment would not significantly affect egg production rates in A. tonsa.