Growth rates of summer nanoplankton (<10 μm) populations in lower Narragansett Bay,Rhode Island,USA

Growth rates of summer (June–September) phytoplankton assemblages and constituent species were measured in 30 diffusion culture experiments. Size-fractionated (<10 m) phytoplankton assemblages were incubated in situ or under simulated in-situ conditions in outdoor tanks connected to a running seawater system. Doubling rates of important species and groups (such as microflagellates) were compared to community biomass doubling rates estimated from ¹⁴C uptake and changes in chlorophyll a concentrations. Division rates of dominant diatom species generally equalled or exceeded community biomass doubling rates, while those of flagellates and non-motile ultraplankters were slower. Maximum division rates of sixteen common diatom species exceeded 2.1 divisions d^-1, while nine had maximum division rates in excess of 3 d^-1. Mean division rates of 12 diatom species exceeded 1 d^-1. Maximum division rates of flagellated species, uncharacterized microflagellates and non-motile ultraplankton assemblages were 2.1, 1.5 and 1.4 d^-1, respectively. Microflagellate and non-motile ultraplankton assemblage doubling rates were less than 0.5 d^-1 in over half of all growth experiments.     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. I. Physiological investigations of Chaetoceros armatum and Asterionella socialis in laboratory cultures

Two species of diatoms, Chaetoceros armatum T. West and Asterionella socialis Lewin and Norris, which account for virtually all of the plant biomass present in the surf zone along the Olympic Peninsula (USA), have been isolated and grown in culture, and comparisons have been made with respect to their physiological behavior in response to aeration, light, temperature, salinity, and nutrients. A. socialis displayed the greater maximum growth rate under most situations, which would give this species a natural advantage over C. armatum. However, in the natural habitat, C. armalum is by far the more important of the two, since the blooms often consist of almost pure stands of this species. This may be due to its apparent ability to use low light levels more efficiently. The maximum growth rate of C. armatum was reached at light levels of 750 to 1000 lux in cultures, whereas A. socialis required a higher light intensity (4000 lux) for maximum growth. C. armatum did not produce its mucilage envelope under any culture conditions. Such a mucilage envelope, with its accompanying clay particles, is invariably present surrounding the diatom chains when they are collected from the natural habitat. The absence of the mucilage envelope (and clay particles) may explain the peculiarities of iron nutrition which were observed in culture experiments, i.e., a dependence on the presence of iron as NaFe-EDTA or ferric citrate, and failure to utilize ferric choloride as a source of Fe for growth.Contribution No. 652 from the Department of Oceanography, University of Washington.     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. IV. Nitrate reductase activity in natural populations and laboratory cultures of Chaetoceros armatum and Asterionella socialis

Nitrate reductase (NR) activity in the surf-zone diatom Chaetoceros armatum T. West was studied in its natural habitat at Copalis Beach, Olympic Peninsula, Washington (USA) over a period of 18 months. Variations of enzyme activity were found, depending on the season and the time of day. General seasonal trends in NR activity were correlated with nitrate concentrations in the surf-zone. The results also indicated that several factors may limit the activity of the enzyme. This is one of the reasons why NR activity could not be correlated with environmental nitrate concentration on the basis of individual measurements at a given point in time. Enzyme inhibition due to the presence of ammonium occurred on several occasions throughout the study period, and could be detected by the NR assay. This phenomenon for the most part masked any pattern of diel periodicity in NR activity that might have been expected to exist in cells in the natural environment. Cultures of C. armatum and Asterionella socialis Lewin and Norris were also studied in the laboratory. A diel fluctuation in NR activity corresponding to the pattern shown in natural phytoplankton by other authors was exhibited in C. armatum under a light-dark regime of 8 h light – 16 h dark and a light intensity of 400 lux.     

Nitrogen regeneration by two surf-zone mysids,Mesopodopsis slabberi andGastrosaccus psammodytes

Nitrogen regeneration by two surf zone mysids,Mesopodopsis slabberi andGastrosaccus psammodytes, was determined under laboratory conditions. The mysids were collected from the lower Sundays River estuary, South Africa, from early spring 1984 to late summer 1985. The forms of nitrogen excreted and the effects of mass, temperature and feeding on excretion rate were determined for each species at three experimental temperatures. Comparison of the forms of nitrogen excreted revealed only slight differences between species, with ammonia the major form and urea and amino acids the secondary excretory products in both cases. Mass significantly influenced the rate of ammonia excretion at all experimental temperatures, with no significant difference in slope (common b=0.602) detected between species. During the day sediment deprivation resulted in a 15% and 20% increase in mean ammonia excretion rates of juvenile and adultG. psammodytes respectively, whereas no significant differences were found at night. The mean ammonia excretion rates of fedM. slabberi andG. psammodytes were 2 and 2.5 times higher than starved excretion rates, respectively.G. psammodytes andM. slabberi recycle 139 to 150 g N per running meter of surf zone per year and 1 007 to 1 208 g N m^-1 yr^-1, respectively. Togehter this constitutes 10% of total phytoplankton nitrogen requirements in the surf zone.     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. X. Chemical composition of the surf diatom Chaetoceros armatum and its major herbivore,the Pacific razor clam Siliqua patula

The surf diatom Chaetoceros armatum T. West, collected from its natural habitat along the Washington coast (USA), had a large inorganic component in the form of a clay coat (consisting of clay minerals, illite and montmorillonite) surrounding the cell chains: 63% of the dry weight of the natural material collected in November was inorganic. The organic fraction was composed of 67.6% lipid, 29.7% protein, and 1.3% carbohydrate. Culture cells were likewise high in protein and lipid and low in carbohydrate. Traces of chitin found in field samples were probably a contaminant, since chitin was absent from cultured cells. This diatom species serves as a major food source throughout the year for the Pacific razor clam Siliqua patula Dixon, which inhabits these same beaches. Protein constituted 47% and lipid 42% of the dry weight of razor clam tissue. The fatty acid distribution in the diatom lipid resembled that previously reported for other diatom species; similarly, the fatty acid distribution of the clam lipid was similar to that previously reported for other bivalve molluscs. The clam fatty acids differed in chain length and degree of saturation from those extracted from its food source, indicating an active fatty acid metabolism in the clam.Contribution No. 995 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Nitrogen regeneration by the surf zone penaeid prawn Macropetasma africanus

Nitrogen excretion of individual Macropetasma africanus (Balss) from an exposed beach/surf zone in Algoa Bay, South Africa was monitored under laboratory and field conditions in relation to body mass, temperature and feeding during 1985. Excretion rate experiments were performed on starved prawns at 15°, 18°, 20° and 25°C, as well as on individuals fed on four different diets (mussel, fish, shrimp and natural diet) at 15° and 20°C. The ratios of the excreted compounds to total nitrogen excreted were similar for the four diets despite differences in their nitrogen content and in the amount of food consumed. At 15° and 20°C, ammonia excretion rates of fed individuals were four to seven times higher than in starved prawns. the excretion rates were not correlated with nitrogen content of diets. M. africanus recycles 1 557 g NH₄–N per metre strip per year or 1 832 g total nitrogen m^-1 yr^-1, which constitute 12 and 14%, respectively, of total phytoplankton requirements of the surf zone. This study indicates that large motile crustaceans, when abundant, can play an important role in nutrient recycling in turbulent marine environments.     

Marine food-web analysis: An experimental study of demersal zooplankton using isotopically labelled prey species

Short-term incubations in seawater containing H¹⁴CO₃ ^- or ³H₂O in place of the naturally predominant isotopes can yield highly radioactive preparations of living phytoplankton or zooplankton. Subsequent in situ incubation of these labelled organisms with the community from which they were taken results in the rapid transfer of radioisotope to those species which prey upon them. This technique has been employed to map a portion of a marine food web involving demersal zooplankton; experiments were conducted in summer and autumn on a coral reef and in a subtropical estuary. Similar results were obtained from these initial experiments at each study site during both seasons. Prey supplied as zooplankton (124 to 410 m nominal diameter), which consisted mainly of Oithona oculata, was fed upon by zooplankton size classes ranging from 410 to 850 m and containing amphipods, ostracods, cumaceans and polychaetes. In experiments employing labelled phytoplankton as prey a wide size spectrum was used (10 to 106 m) in order to include representative samples of most of the available planktonic autotrophs as estimated by primary production measurements. In two separate experiments, only 7 out of 63 samples evidenced grazing of phytoplankton by demersal zooplankters. In contrast, labelled diatom auxospores, employed in one experiment as they constituted the most numerically abundant species in the water column, were found to be grazed upon in nearly half the samples examined.     

Seasonal variations in the utilization of ammonium and nitrate by photoplankton in Vineyard Sound,Massachusetts, USA

The nitrogenous nutrition of the phytoplankton in Vineyeard Sound, Massachusetts, USA was investigated over a 15-month period. Highest rates of ammonium uptake were observed immediately prior to, or during, the diatom bloom periods, and with one exception were found in the <10 m size class. The saturating rate of ammonium uptake correlated well with temperature and gave Q₁₀ values of 2.6–3.2; correlations with ambient solar irradiation were not nearly so clear. Uptake rates of ammonium exceeded those of nitrate except during the winter bloom of the diatom Rhizosolenia delicatula; yet calculation of the f ratio revealed that nitrate was relatively important in the nitrogenous nutrition of the phytoplankton throughout the year.Contribution no. 5096 from the Woods Hole Oceanographic Institution     

Spring and summer growth rates of subarctic Pacific phytoplankton assemblages determined from carbon uptake and cell volumes estimated using epifluorescence microscopy

In order to determine whether phytoplankton growth rates were normal or depressed, total plant carbon (g l^–1) and in situ production rates (g C l^–1 d^–1) were measured for phytoplankton assemblages at Weathership Station P (50°N; 145°W) and at 53°N; 145°W in the subarctic Pacific in May and August 1984. Plant carbon, estimated from cell volumes determined using epifluorescence microscopy, was distributed as follow: 28% in the <2 m fraction, 38% in the 2 to 5 m size fraction, and the remainder in size classes >5 m. Carbon-specific growth rates (k), as doublings d^–1, were calculated for the phytoplankton assemblages as a whole at each sampling depth down to 100 m for three days in May and for four days in August. The populations in the upper part of the euphotic zone showed average doubling rates of 1 d^–1 and thus appeared to be growing at rates normally expected for the prevailing conditions of light and temperature. The low chlorophyll concentrations (0.3 to 0.4 mg chl a m^–3) characteristically found in this oceanic region do not seem to be due to very slow growth of algal populations.Contribution No. 1695 of the School of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Abundance and production rates of floating diatom mats (Rhizosolenia castracanei and R. imbricata var. shrubsolei) in the Eastern Pacific Ocean

Abundance and production rates were measured on freefloating mats composed of the diatoms Rhizosolenia castracanei and R. imbricata var. shrubsolei in the California Current and boundary waters of the central North Pacific during October, 1980. Mats ranged from 3.0 to 10.6 cm in maximum length and had a mean volume of 6.4 ml. Production rates of the diatom mats averaged 4.0 g C colony^-1 h^-1, more than 10³ times higher than that of an equal volume of surrounding water. However, because of their low density at these sites, about 1 mat m^-3, diatom mats contributed only about 1% of the total primary production. The two large Rhizosolenia species comprised almost all of the phytoplankton biomass within the mats. Rhizosolenia species rarely occurred in seawater between mats, where the phytoplankton community was dominated by a diverse array of nannoplankton. The Rhizosolenia species in the mats appeared to be in healthy condition and contained intracellular bacteria. The very high production rates of the colonies indicate high nutrient demand and, since these particular diatoms lack the symbiotic cyanobacterium Richelia intracellularis, which is implicated in N-fixation in other species of Rhizosolenia, our results indicate that other nutrient sources must be present. We discuss the potential role of the intracellular bacteria in nitrogen fixation.     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. XI. Regeneration of ammonium in the surf environment by the Pacific razor clam Siliqua patula

A 20 month field study was conducted on ammonium excretion rates of the Pacific razor clam Siliqua patula Dixon along the beaches of Washington State, USA. Excretion rates of all those nutrients likely to be regenerated in sufficient quantity to affect surf diatom growth were measured; ammonium appeared to be the most important metabolite. Excretion of ammonium by razor clams far exceeded that by other beach fauna. Ammonium excretion rates of razor clams were positively correlated with shell length, but no correlation between ammonium excretion rate and water temperature was evident. This may be an artifact or may represent some degree of seasonal acclimation of the species to temperature. Weight-specific ammonium excretion rates were negatively related to clam size, indicating a possible large (and unknown) contribution of regenerated ammonium by smaller clams in their first year of growth; smaller clams were rarely captured during this study.Contribution No. 1048 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Availability of colloidal ferric oxides to coastal marine phytoplankton

Cell growth of a coastal marine diatom, Phaeodactylum tricornutum (stock cultures), and two red tide marine flagellates, Heterosigma akashiwo and Gymnodinium mikimotoi (stock cultures), in the presence of soluble chelated Fe(III)-EDTA (1:2) and of four different phases of ferric oxide colloids were experimentally measured in culture experiments at 20°C under 3000 lux fluorescent light. Soluble Fe(III)-EDTA induced the maximal growth rates and cell yields. The short-term uptake rate of iron by H. akashiwo in Fe(III)-EDTA medium was about eight times faster than that in solid amorphous hydrous ferric oxide (Fe₂O₃·xH₂O) medium. In culture experiments supplied with four different ferric oxide forms, the orders of cell yields are amorphous hydrous ferric oxide>-FeOOH (lepidocrocite)>Fe₅O₇(OH)·4H₂O (hydrated ferric oxyhydroxide polymer >-FeOOH (goethite). The specific growth rates () at logarithmic growth phase in Fe(III)-EDTA, amorphous hydrous ferric oxide and -FeOOH media were significantly greater than those in Fe₅O₇ (OH)·4H₂O and -FeOOH media. The thermodynamically stable forms such as Fe₅O₇(OH)·4H₂O and -FeOOH supported a little or no phytoplankton growth. The iron solublities and/or proton-promoted iron dissolution rates of these colloidal ferric oxides in seawater at 20°C were determined by simple filtration techniques involving -activity measurements of ⁵⁹Fe. The orders of solubilities and estimated dissolution rate constants of these ferric oxides in seawater were consistent with that of cell yields in the culture experiments. These results suggest that the availability of colloidal iron to provide a source of iron for phytoplankton is related to the thermodynamic stability and kinetic lability of the colloidal ferric oxide phases, which probably control the uptake rate of iron by phytoplankton.     

Interactive effects of nitrogen and dissolved inorganic carbon on photosynthesis,growth, and ammonium uptake of the macroalgae Cladophora vagabunda and Gracilaria tikvahiae

Dissolved inorganic carbon (DIC) is rarely considered limiting for macroalgae, but some research suggests that under conditions of N sufficiency, photosynthetic capacity is enhanced with DIC enrichment. During spring (April–May) and summer (July–August) 1993, we investigated the interactive effects of nitrogen (N) and DIC on photosynthetic capacity, growth, and nutrient uptake rates of the macroalgae, Cladophora vagabunda (L.) van den Hoek and Gracilaria tikvahiae (McLachlan), dominant species in a temperate eutrophic estuary (Cape Cod, Massachusetts, USA). Water-column CO₂ concentrations showed significant diurnal fluctuations, ranging from a morning CO₂ peak (21 M) to an afternoon low (13 M) during summer, probably associated with metabolic activities in a thick algal mat. Results from instantaneous photosynthesis measurements and microcosm experiments indicate that DIC limits photosynthetic capacity and growth rates of C. vagabunda during summer, perhaps related to tissue N sufficiency and low water-column CO₂ concentrations. For example, this species showed enhanced growth (F=8.69, P<0.02) under DIC but not N enrichment. G. tikvahiae showed marginal DIC enhancement of maximum photosynthetic rate, while growth was significantly stimulated by addition of N. Reduced thallus N of this species during the summer further identifies N as the primary factor limiting growth. In addition, G. tikvahiae has the ability to use DIC in its several forms, while C. vagabunda primarily uses dissolved CO₂. DIC enrichment resulted in a depression of NH₄ ⁺ uptake rates for both species, particularly during summer at saturating (60 M) ammonium levels, suggesting competition between NH₄ ⁺ uptake and DIC acquisition under conditions of N sufficiency. Dominance of C. vagabunda and G. tikvahiae in areas undergoing eutrophication has been attributed to their successful procurement and storage of N as well as to high growth rates. The present study revealed that under conditions of N sufficiency during summer, DIC may control rates of production of these opportunistic macroalgae.     

Life history of the sandy-beach amphipod Dogielinotus loquax (Crustacea: Dogielinotidae) from the outer coast of Washington,USA

Dogielinotus loquax Barnard, a common intertidal macroinvertebrate on exposed ocean beaches near Grays Harbor, Washington, USA, is distributed from mid to mean high tidal levels, with a mean density of 1 830 m^-2 in 1975–1976. Location of maximum density within the intertidal zone is related to surf intensity and varies seasonally. The species is iteroparous and has an average sex ratio of 1:1. Temperature constraints on growth and egg development rates apparently influence the timing of reproduction. Two dominant recruitments occur per year, one in early spring (the summer generation) and another in late summer (the overwintering generation). The sampling design and density estimates permitted estimation of the mortality rate for the summer generation. Shorebird predation is suspected to affect summer generation abundance. Dogielinotus loquax will soon be elevated to the type of a new genus, Proboscinotus (Bousfield and Tzvetkova, in press)Contribution no. 1271 from the Department of Oceanography, University of Washington; Seattle, Washington 98195, USA     

Rates of primary productivity and growth in<Emphasis Type="Italic"> Ecklonia radiata</Emphasis> measured at different depths,over an annual cycle,at West Island,South Australia

The pattern of growth (biomass accumulation) in Ecklonia radiata throughout the year and across a depth profile was investigated using the traditional hole-punch method, and the information presented in context with concurrently measured in situ net productivity rates. The rate of net daily productivity showed a lack of consistent seasonal variability, remaining constant throughout the year at two of the four depths measured (3 m and 12 m), and becoming higher during winter at another (5 m). Throughout the year, rates of net daily productivity differed significantly across the depth profile. Net daily productivity rates averaged 0.017 g C g^–1 dwt day^–1 and 0.005 g C g^–1 dwt day^–1 at a depth of 3 m (1,394 mol O₂ g^–1 dwt day^–1) and 10 m (382 mol O₂ g^–1 dwt day^–1) respectively. In contrast, the biomass accumulation rate of E. radiata was highly seasonal, with low rates of growth occurring in autumn (0.002 g dwt g^–1 dwt day^–1 at both 3 and 10 m) and summer (0.007 and 0.004 g dwt g^–1 dwt day^–1 at 3 and 10 m respectively) and higher rates in spring (0.016 and 0.007 g dwt g^–1 dwt day^–1 at 3 and 10 m respectively) and winter (0.015 and 0.008 g dwt g^–1 dwt day^–1 at 3 and 10 m respectively). The proportion of assimilated carbon used for biomass accumulation varied throughout the year, between 5% and 41% at 3 m and between 28% and 128% at 10 m. The rates of biomass accumulation at all depths represented only a small proportion of the amount of carbon assimilated annually.Communicated by P.W. Sammarco, Chauvin     

Carbon fixation in marine phytoplankton: carboxylase activities and stable carbon-isotope ratios; physiological and paleoclimatological aspects

We measured the activity of three carboxylases: RuBP carboxylase, PEP carboxylase and PEP carboxykinase of marine phytoplankton species in culture and in natural communities. Activities of the three carboxylases were measured simultaneously with stable carbon-isotope ratios. The enzymatic activities have been used to estimate the importance of carboxylation and its impact on the ¹³C:¹²C ratio (expressed as ¹³C). The marine phytoplankton species in culture were Fragilariopsis kerguelensis, Nitzschia turgiduloides, Skeletonema costatum, Phaeodactylum tricornutum, Isochrysis galbana, Dunaliella marina, and Prorocentrum micans, and the field samples were collected from different depths off the coast of Portugal (August/September, 1981). Our results indicate that, as in terrestrial plants, the ¹³C value is a good indicator of the extent of carboxylation. RuBP carboxylase activity was always predominant, whereas the ¹³C value never reached values typical of the C₄ pathway. The carboxylases could be PEP carboxylase (in dinoflagellates) or PEP carboxykinase (in diatoms). carboxylation increased at the end of the exponential growth phase in a diatom culture and with increased biomass in natural samples. We interpret these increases as an adaptative response mechanism to poor environmental conditions, especially to low light intensity.     

Seasonal growth in Laminaria longicruris: Relations with dissolved inorganic nutrients and internal reserves of nitrogen

Observations have been made on seasonal fluctuations in dissolved inorganic nutrients, internal reserves of nitrogen and growth rates in Laminaria longicruris. The onset of winter growth in shallow-water stations (6 and 9 m) correlated well with improved dissolved nitrate conditions in the sea. During the winter, reserves of NO ₃ ^- were accumulated by the plants and reached maximum values of 150 moles per g fresh weight in March. This represents a concentration factor of approximately 28,000 over the ambient levels, or an internal nitrogen reserve of 2.1% of the dry weight of the tissue. Depletion of this nitrogen pool followed the disappearance of the external NO ₃ ^- with a lag period of up to 2 months. Rapid kelp growth was measured during this period. Reserves of organic nitrogen also reached maximum values in March and declined slowly throughout the summer into autumn. It is suggested that the combined inorganic and organic nitrogen reserves sustain the rapid growth rates into July and at reduced rate through the late summer. Fertilization of an experimental perimental kelp bed with NaNO₃ increased the internal plant reserves of NO ₃ ^- and produced a much improved summer growth rate. The enriched plants developed very small reserves of carbohydrate during the rapid summer growth phase.NRCC No. 15549.     

Effects of food quality on the reproductive success of Acartia tonsa and Acartia hudsonica: laboratory observations

The effect of the chemical composition of food on the reproductive success of the copepods Acartia tonsa Dana and A. hudsonica Pinhey was studied in the laboratory. Laboratory-reared individuals were fed one of three monoalgal diets at different stages of growth: the diatom Thalassiosira weissflogii, the flagellate Rhodomonas lens and the dinoflagellte Prorocentrum minimum. The diet was analyzed for carbon, nitrogen, protein, carbohydrate and fatty acid content. Reproductive success was measured as eggs female^-1 day^-1 (E _r) and as the hatching success of the eggs. The E _r of Acartia spp. was correlated with protein and specific fatty acids [16:17 (negative), 20:53, 22:63, and 18:0 (positive)] and, especially, the fatty acid composition of the algae expressed as the 3:6 and 20:22 fatty acid ratios. The youngest diatom cultures and exponentially-growing flagellates displayed the highest E _r; the lowest E _r was recorded for females fed the senescent diatom cultures. The development time of eggs was affected by the age of the phytoplankton culture fed to the female. Hatching success of eggs decreased with the age of the algal culture, but no correlation was found with the meansured chemical components of the food.     

Omnivorous feeding behavior of the Antarctic krill Euphausia superba

Feeding experiments were conducted at Palmer Station from December 1985 to February 1986 to examine the potential role of copepod prey as an alternative food source for Euphausia superba. Copepod concentration, copepod size, phytoplankton concentration, the duration of krill starvation and the volume of experimental vessels were altered to determine effects on ingestion and clearance rates. Krill allowed to feed on phytoplankton and copepods in 50-litre tubs showed greatly increased feeding rates relative to animals feeding in the much smaller volumes of water traditionally used for krill-feeding studies. Clearance rates on copepods remained constant over the range of concentrations offered, but clearance rates on phytoplankton increased linearly with phytoplankton concentration. Feeding rates increased when larger copepods were offered and when krill were starved for two weeks prior to experiments. Clearance rates of krill feeding on copepods were higher than, but not correlated with, their clearance rates on phytoplankton in the same vessel. E. superba may have a distinct mechanism for capturing copepods, perhaps through mechanoreception. Although our observed clearance rate of 1055 ml krill^-1 h^-1 indicates that krill can feed very efficiently on copepod prey, such feeding would meet less than 10% of their minimum metabolic requirements at the typical copepod concentrations reported for Antarctic waters. However, substantial energy could be gained if krill fed on the patches of high copepod concentrations occasionally reported during the austral summer, or if krill and copepods were concentrated beneath the sea ice during the winter or spring months. Our results, indicating efficient feeding on zooplankton and higher clearance rates on phytoplankton than previously believed, represent a step towards balancing the energy budget of E. superba in Antarctic waters.     

Production and respiration of overall plankton and ultraplankton communities at the entrance to the Gulf of Finland in the Baltic Sea

Primary productivity and respiration of the overall plankton community and of ultraplankton (organisms passing through a 3-m Nuclepore filter) were studied at the entrance to the Gulf of Finland during the growth season in 1982. Data of the respiration measurements from previous years are also presented. During the development of a diatom spring bloom, the algal component could be successfully separated from the bacterial component by size fractionation with a 3-m Nuclepore filter and thus the algal respiration could be approximated, being on the order of 10 to 20% of the gross production. After the phytoplankton spring maximum, bacteria played an important role in mediating the energy flow from phytoplankton exudates to higher trophic levels. Maximum values of 1 230 and 740 mg O₂ m^-2 d^-1 were recorded for overall and for ultraplankton respiration, respectively, during late July. High productivity values coupled with low phytoplankton biomass and low inorganic nutrient values were also recorded in late July, indicating effective nutrient regeneration and rapid turnover of the plankton community. During late summer, a considerable fraction (over 30%) of phytoplankton production was released as exudates, suggesting that much of the energy is channeled to higher trophic levels via bacterial pathways rather than by direct herbivorous grazing during this season. The summer development of phytoplankton community structure and functioning is strongly controlled by hydrographic conditions, i.e. by nutrient inputs via upwelling and by water temperature. A carbon budget for late summer indicated that bacteria may contribute only up to 50% of the overall respiration of the plankton community, which suggests that heterotrophs other than bacteria play an important role in nutrient regeneration. The present study stresses the importance of energy flow via the phytoplankton exudatebacteria-micrograzer pathway in relatively oligotrophic, brackish water ecosystems.