Ecotypic differentiation of Laminaria longicruris in relation to seawater nitrate concentration

The ultraplankton (cell diameters >3 μm), which compromises about 70% of the biomass of phytoplankton in subtropical surface waters near Oahu, Hawaii, was isolated for growth rate studies. The specific growth rate (μ) was estimated from the rate of increase of the chlorophyll biomass during incubations in the absence of grazers. This growth rate of the ultraplankton ranged from 0.037 to 0.071 h^?1 (=1.3 to 2.5 doublings d^?1) during a period when P:B ratios of 5 to 14.5 μg C μg^?1 chl a h^?1 prevailed. The co-occurrence of atypically high P:B ratios and nonlimiting ambient nutrient concentrations suggests that the calculated values are higher than those characteristic of such subtropical ecosystems in general. Rates of ammonium uptake and photosynthesis by the >3 μm fraction were also compared to those of larger fractions. Organisms in the >3 μm fraction assimilated NH ₄ ⁺ at a rate which was about 75% greater than that of the 3 to 20 μm size fraction. Comparison of μ and P:B data collected over a 2 mo period (November–December, 1980) shows that the correlation between these two rate indices is nonlinear. The predominance of small-celled phytoplankton in oligotrophic waters is explained, in part, by its higher μ, its higher nutrient assimilation rates, and the absence of its loss through sedimentation.     

Phytoplankton exudate release as an energy source for the growth of pelagic bacteria

The release of dissolved organic carbon (DOC) from phytolankton during photosynthesis, and the utilization of this carbon by planktonic bacteria, was studied using ¹⁴CO₂ and selective filtration. Natural sea water samples from a coastal area of the Northern Baltic Sea were incubated in the laboratory for detailed studies, and in situ for estimation of annual dynamics. In a laboratory incubation (at +1°C) the concentration of ¹⁴C-labelled dissolved organic carbon increased for about 2 h and then reached a steady state, representing about 0. 1% of the total DOC. Labelled organic carbon in the phytoplankton and bacterial fractions continued to increase almost linearly. The continuous increase in the bacterial fraction is thought to represent almost instantaneous utilization of the DOC released from the phytoplankton during photosynthesis. As an annual average, in 4 h in situ incubations, about 65% of the labelled organic carbon was found in the phytoplankton fraction (>3 m), about 27% in the bacterial fraction (0.2 to 3 m) and the remaining 8% as DOC (<0.2 m). Large variations in these percentages were recorded. The measured annual primary production was 93 g C m^-2 (March to December), and the estimated bacterial production due to phytoplankton exudates 29 g C m^-2. This represents a release of DOC of about 45% of the corrected annual primary production of 110 g C m^-2 (assuming a bacterial growth efficiency of 0.6).     

Photosynthetic characteristics of nanoplankton and picoplankton from the surface mixed layer

Nanoplankton and picoplankton primary production has been studied at two oceanic stations in the Porcupine Sea-bight and at one shelf station in the Celtic Sea. At both sites, low wind conditions in June and July 1985 resulted in greatly reduced vertical turbulent mixing and a secondary, temporary thermocline developed in what is usually a well-mixed surface layer; as a result, there was physical separation of the phytoplankton within two zones of the surface mixed layer. The photosynthetic characteristics of three size fractions (>5 m, <5 to >1 m and <1 to >0.2 m) of phytoplankton populations from the two zones have been measured. Phytoplankton was more abundant at the oceanic stations and chlorophyll a values were between 1.3 and 2.2 mg chlorophyll a m^-3, compared with 0.3 to 0.6 mg chlorophyll a m^-3 at the shelf station; at both stations, numbers of cyanobacteria were slightly higher in the lower zone of the surface mixed layer. There was no effect of the temporary thermocline on the vertical profiles of primary production and most phtosynthesis occurred in the surface 10 m. Photosynthetic parameters of the three size fractions of phytoplankton have been determined; there was considerable day-to-day variation in the measured photosynthetic parameters. Assimilation number (P _m ^B ) of all >5 m phytoplankton was lower for the deeper than for the surface populations, but there was little change in initial slope (a ^B ). The small oceanic nanoplankton (<5 to >1 m) showed changes similar to the >5 m phytoplankton, but the same size fraction from the shelf station showed changes that were more like those shown by the picoplankton (<1 m) viz, little change in P _m ^B but an increase in a ^B with depth. Values of a ^B were generally greater for the picoplankton fraction than for the larger phytoplankton, but values of adaptation parameter (I _k )(=P _m ^B /) were not always less. There was little evidence to support the hypothesis that these populations of picoplankton were significantly more adapted to low light conditions than the larger phytoplankton cells. When photosynthetic parameters of the picoplankton were normalised to cell number (P _m ^C /a ^C ) rather than chlorophyll a, P _m ^C was comparable to other published data for picoplankton, but a ^C was much lower. The maximum doubling time of the picoplankton at saturating irradiance is calculated to be ca. 8.5 h for the oceanic population and ca. 6.2 h for the shelf population.     

Size structure of phytoplankton biomass and photosynthesis in subtropical Hawaiian waters

In a subtropical Hawaiian ecosystem, phytoplanton size structure analyses (November–December, 1980) showed that ultraplankton (>3μm), nanoplankton (>20μm) and netplankton (>20μm) accounted for ca. 80, 98, and 2% of total chlorophyll standing stock, respectively, on the basis of chlorophyll. Similar trends were evident, for other biomass indices (e.g. cell numbers, total cell volume, ATP, particulate organic carbon, particulate organic nitrogen). The ultraplankton fraction consisted primarily of small flagellates (1 to 3 μm diam) and coccoid cells (?1 μm diam); the 3 to 20 μm fraction was represented by dinoflagellates, coccolithophores, diatoms, and chrysophytes; and the netplankton fraction consisted principally of dinoflagellates and centric diatoms. Community photosynthesis had a size distribution similar to that of biomass. Sinking rates for the 3 μm, 3 to 20 μm, and >20 μm fractions averaged 0.0, 0.09, and 0.29m d^?1, respectively. The absence of measurable sinking rates for the ultraplankton, together with the relative abundance of biomass in this fraction, result in very small phytoplankton losses due to sinking in such subtropical surface waters.     

Fractionated phytoplankton primary production,exudate release and bacterial production in a Baltic eutrophication gradient

The distribution of phytoplankton primary production into four size fractions (>10 m, 10-3 m, 3-0.2 m and <0.2 m), the utilization of algal exudates by bacteria and the bacterial production were studied in a eutrophication gradient in the northern Baltic proper. The polluted area exhibits substantially increased nutrient, especially nitrogen, levels while only minor differences occur in salinity and temperature regimes. Total primary production was 160 g C · m^-2 · yr^-1 at the control station and about 275 g C · m^-2 · yr^-1 at the eutrophicated stations. The estimated total exudate release was 16% of the totally fixed ¹⁴CO₂ in the control area and 12% in the eutrophicated area (including the estimated bacterial uptake of exudates). The difference in¹⁴CO₂ uptake rates between incubation of previously filtered water (<3, <2, <1 m) and unfiltered water was used to estimate bacterial uptake of phytoplankton exudates which were found to contribute about half of the estimated bacterial carbon requirement in both areas. Bacterial production was estimated by the frequency of dividing cells (FDC) method as being 38 g C · m^-2 · yr^-1 at the control station and 50 g C · m^-2 · yr^-1 at the eutrophicated stations. To estimate the mean in situ bacterial cell volume a correlation between FDC and cell volume was used. The increased annual primary production in the eutrophicated area was due mainly to higher production during spring and autumn, largely by phytoplankton cells (mainly diatoms) retained by a 10 m filter. Primary production duringsummer was similarin the two areas, as was the distribution on different size fractions. This could possibly explain the similar bacterial production in the trophic layers at all stations since the bulk of bacterial production occurs during summer. It was demonstrated that selective filtration does not quantitatively separate photoautotrophs and bacteria. A substantial fraction of the primary production occurs in the size fraction <3 m. The primary production encountered in the 3-0.2 m fraction was due to abundant picoplankton (0.5 to 8 · 10⁷ ind · l^-1), easily passing a 3 m filter. The picoplankton was estimated to constitute up to 25% of the total phytoplankton biomass in the control area and up to 10% in the eutrophicated area.     

Physiological ecology of picoplankton in the North Sea

The distribution of cyanobacteria in the surface waters of the North Sea was measured during July 1987. Numbers of cyanobacteria ranged from 2.5x10⁶ to 1.7x10⁸ cells 1^-1. In the majority of stations, cyanobacterial numbers were highest in the near-surface water and a subsurface maximum was found at only one station. The distribution of ¹⁴C among the end-products of photosynthesis was determined for picoplankton (<1 m) and other phytoplankton >1 m throughout the North Sea. The majority of label was found in the protein fraction of both picoplankton and >1 m phytoplankton; incorporation into lipids and polysaccharides plus nucleic acids was much lower. We interpret the large incorporation into protein to be a consequence of nutrient limitation of these natural assemblages. Photosynthetic parameters of the two size fractions were also determined. Assimilation number (P _m ^B ) and initial slope were greater for the picoplankton fraction than for phytoplankton >1 m but there was no evidence of significant photoinhibition of either fraction at irradiances up to 1 000 E m^-2 s^-1.     

Response of the microbial food web to manipulation of nutrients and grazers in the oligotrophic Gulf of Aqaba and northern Red Sea

The control mechanisms within the pelagic microbial food web of the oligotrophic Gulf of Aqaba and the northern Red Sea were investigated in the spring of 1999. Nutrient conditions and potential grazer impact were manipulated in a series of dilution experiments. Ambient nutrient concentrations and autotrophic biomass were very low (0.23–1.21 µmol NO₃ l^–1, 0.06–0.98 µmol NH₄ l^–1, 1.08–1.17 µmol Si l^–1, 0.08–0.12 µmol P l^–1, 0.15–0.36 µg chlorophyll a l^–1). The planktonic community was characterized by low abundances [3.0–5.5×10⁵ heterotrophic bacteria ml^–1, 0.58–7.2×10³ ultraphytoplankton <8 µm ml^–1 (small eukaryotic photoautotrophs and Prochlorococcus sp., excluding Synechococcus sp.), 0.45–4.4×10⁴ Synechococcus sp. ml^–1, 0.32–1.2×10³ heterotrophic nanoflagellates ml^–1, 1.3–3.8×10³ phytoplankton >8 µm l^–1, 0.93–5.4×10² microzooplankton l^–1] and dominated by small forms (0.2–8 µm). Dinoflagellates and oligotrichous ciliates were the most common groups in initial samples among the phytoplankton >8 µm and microzooplankton, respectively. Results show that bottom-up and top-down control mechanisms operated simultaneously. Small organisms were vulnerable to grazing, with maximum grazing rates of 1.1 day^–1 on heterotrophic bacteria and 1.3 day^–1 on ultraphytoplankton. In contrast, algae >8 µm showed stronger signs of nutrient limitation, especially when the final assemblages were dominated by diatoms. Synechococcus sp. were not grazed and only showed moderate to no response to nutrient additions. The high spatial and temporal variation of our results indicates that the composition of the planktonic community determines the prevailing control mechanisms. It further implies that, at this transitional time of the year (onset of summer stratification), the populations fluctuate about an equilibrium between growth and grazing.Communicated by O. Kinne, Oldendorf/Luhe     

Zooplankton abundance and grazing at Davies Reef,Great Barrier Reef,Australia

Zooplankton abundance and grazing on autotrophic and heterotrophic particulate matter were measured along a transect across Davis Reef (18°5S; 147°39E) and in the back-reef lagoon over tidal and diel cycles during austral winter (August 1984). Zooplankton entering the reef from the surrounding shelf waters decreased in abundance over the reef flat, presumably because of predation. Within the reef lagoon, maximum daytime densities of pelagic copepods occurred during high water, suggesting an external input. At night, water-column zooplankton biomass increased by a factor of 2 to 3 due to the emergence of demersal reef zooplankton. Zooplankton grazing rates on heterotrophic particulate matter (bacteria + detritus and Protozoa) compared to phytoplankton were higher on the reef flat than on the fore-reef or lagoon. Within the lagoon, zooplankton grazing rates on heterotrophic material were maximum during high water, coincident with maximum tidal concentrations of particulate organic carbon. The combined demersal and pelagic zooplankton community were often able to crop 30% of the daily primary production by >2µm phytoplankton. However, >50% of phytoplankton biomass was in cells <2µm, presumably unavailable to these zooplankton. Our particulate production and ingestion measurements, together with zooplankton carbon demand extrapolated from respiration estimates, suggest that the zooplankton community of Davies Reef derives much of its nutrition from detritus.Joint contribution from the University of Maryland, Center for Environmental and Estuarine Studies (No. 2015), and the Microbial Ecology on a Coral Reef Workshop (MECOR No. 19)     

Some experiments on phosphate assimilation by coastal marine plankton

A study of phosphate assimilation by coastal marine plankton revealed that both phytoplankton and microheterotrophs incorporated radioactive phosphorus (³³P). Size fractionation of the particulate matter (using 1 m pore diameter Nucleopore^® membrane filters), antibiotic treatment (using garamycin), and independent estimaties of photoautotrophic (¹⁴CO₂ uptake) and heterotrophic (³H-glucose uptake) activities were employed to separate phyto- and bacterioplankton phosphate uptake. Results indicated that phytoplankton ³³P-uptake was best estimated by the fraction of particulate matter retained on the 1 m membrane filters. Usually, less than 10% of the phytoplankton (based on chlorophyll a measurements) passed the 1 m pore-diameter filters, whereas about 90% of the heterotrophic activity passed. At least 50% of the ³³P-uptake was associated with the <1 m fraction. It may be possible to resolve the phytoplankton and bacterial contributions to ³³P-uptake by comparing the percent of total ³³P-uptake with the percent of total ³H-glucose uptake associated with the >1 m fraction.     

Seasonal variations in microzooplankton grazing in the region of the Subtropical Convergence

Microzooplankton grazing and community structure were investigated in the region of the Subtropical Convergence (STC) during three cruises of the South African Antarctic Marine Ecosystem Study (SAAMES) in austral summer (January/February 1993; December 1994/January 1995) and winter (June/July 1993). Chlorophyll a concentrations were consistently dominated by the <20 m size fraction during all three cruises, while the contribution of the microphytoplankton (>20 m) to total chlorophyll a concentrations varied considerably between cruises. Microzooplankton communities were numerically dominated by protozoans comprising ciliates (aloricates and tintinnids) and dinoflagellates. Instantaneous growth coefficients of phytoplankton in the vicinity of the STC showed no seasonal trends. However, marked seasonal differences were observed in the size structure of the phytoplankton. The grazing impact of microzooplankton was highest when the <20 m chlorophyll fraction contributed >95% of the total. Under these conditions, the instantaneous grazing rates ranged between 0.15 and 0.66 d^-1. These correspond to daily losses of 14 to 48% of the inntial standing stock and between 45 and 81% of the potential primary production. At stations where microphytoplankton contributed significantly (-20%) to total chlorophyll concentrations, the grazing coefficients were lower, ranging between 0 and 0.53 d^-1. This corresponds to a loss of <41% of the initial standing stock, or between 0 and 56% of the potential production. Our data suggest that microzooplankton represent the main grazing sink for production when the <20 m chlorophyll size-class dominates total chlorophyll. These facts suggest that the efficiency of the biological pump may vary over time.     

Microzooplankton herbivory on the Grand Bank (Newfoundland,Canada): A seasonal study

Grazing impact of microzooplankton on phytoplankton was investigated on the Grand Bank, Newfoundland, Canada, in April, July and October 1984, using a seawater dilution method. In April a large proportion of chlorophylla was in the microplankton size fraction (> 20µm) while in mid-summer and fall most was in the nanoplankton size fraction (< 20µm). Diatoms were the dominant phytoplankters in April, while undetermined flagellates and coccolithophores were abundant in other seasons. Major grazers were oligotrichous ciliates in all seasons. Instantaneous grazing rates on nanophytoplankton, as measured by changes in chlorophylla, varied from 0.12 to 0.43 d^–1 and those on microphytoplankton from 0.19 to 0.68 d^–1. Grazing rates did not change over 24 and 48 h intervals. This level of grazing corresponded to a daily loss of about 20 and 30% of standing stock of chlorophylla and about 50 and 70% loss of potential production in the two size fractions respectively. Taxon-specific grazing rates, calculated from microscopic enumeration, showed that small diatoms were grazed heavily, and their growth was controlled by grazing in late spring. In late summer and fall, undetermined flagellates and coccolithophores were also grazed at high rates but their growth rates were higher than the grazing rates, and therefore, were not controlled by microzooplankton. In general, microzooplankton grazed on whatever appropriate sized food was dominant in the experimental water. Their potential ability to control the growth of certain food species may be one of the causes determining the species composition of phytoplankton communities.     

Distribution and structure of heterotrophic protist communities in the northeast equatorial Pacific Ocean

The distribution and structure of heterotrophic protist communities and size-fractionated chlorophyll a were studied during the Korea Deep Ocean Study 98 (KODOS 98) research expedition (July 1998) in the northeast equatorial Pacific Ocean (5–11°N). Areas of convergence and divergence formed at the boundaries of the South Equatorial Current (SEC), North Equatorial Current (NEC), and North Equatorial Counter Current (NECC) during the expedition. Water column physicochemical characteristics significantly influenced the size structure of heterotrophic protist communities. Intense vertical mixing and high nutrient and chlorophyll a concentrations characterized SEC and NECC areas, which were affected by converging and diverging water masses, respectively. Nanophytoplankton dominated in SEC and NECC areas; both areas also had relatively high heterotrophic protist biomasses (average 743 µg C m^–2). NEC areas were characterized by a stratified vertical structure, low nutrient and chlorophyll a concentrations, and picophytoplankton dominance. The heterotrophic protist biomass in NEC areas averaged 414 µg C m^–2; nanoprotists (<20 µm) dominated the community. The nanoprotist biomass comprised 49–54% of the total heterotrophic protist biomass in SEC/NECC areas and 67–72% in NEC areas. The biomass of heterotrophic protists was higher in SEC/NECC areas than in NEC areas, but the relative importance of nanoprotists was greater in NEC areas than in SEC/NECC areas. Heterotrophic dinoflagellates were dominant components of the <20 µm and >20 µm size classes in both water columns. The biomass of heterotrophic protists significantly correlated with the net-, nano-, and picophytoplankton biomass in SEC/NECC areas and with the nano- and picophytoplankton biomass in NEC areas. Heterotrophic protists and phytoplankton also showed strong positive correlation in the study area. The size structure of the phytoplankton biomass coincided with that of heterotrophic protists; the heterotrophic protist biomass positively correlated with the protists prey source. These relationships suggest that the community structure of heterotrophic protists and the microbial food web depended on size classes within the phytoplankton biomass. Microzooplankton grazing and phytoplankton growth rates were higher in SEC/NECC areas than in NEC areas. In contrast, the potential primary production grazed by microzooplankton was relatively high in NEC areas (127.3%) compared with SEC/NECC areas (94.6%). Our results indicate that the relative importance and size structure of heterotrophic protists might vary according to two distinct water column structures.Communicated by T. Ikeda, Hakodate     

Trophodynamic function of copepods,appendicularians and protozooplankton in the late summer zooplankton community in the Skagerrak

The study was carried out in the Skagerrak during late summer when population development in the pelagic cycle culminated in the yearly maximum in zooplankton biomass. The cyclonic circulation of surface water masses created the characteristic dome-shaped pycnocline across the Skagerrak. The large dinoflagellate Ceratium furca dominated the phytoplankton biomass. Ciliates and heterotrophic dinoflagellates were the major grazers and, potentially, consumed 43–166% of daily primary production. The grazing impact of copepods was estimated from specific egg production rates and grazing experiments. The degree of herbivory differed between species (14–85%), but coprophagy (e.g. feeding on fecal pellets) and ingestion of microzooplankton were also important. The appendicularian Oikopleura dioica was present in lower numbers than copepods, but cleared a large volume of water. The grazing impact of copepods and O. dioica was estimated to 57±24% and 12±12% of daily primary production, respectively. Sedimentation of organic material (30 m) varied between 169 and 708 mg C m^–2 day^–1, and the contribution from the mesozooplankton (copepod fecal pellets and mucus houses with attached phytodetritus of O. dioica) was 5–33% of this sedimentation. Recycling of fecal pellets and mucus houses in the euphotic zone was 59% and 36%, respectively. However, there was a high respiration of organic material by microorganisms in the mid-water column, and 34% of the sedimenting material actually reached the benthic community in the deep, central part of the Skagerrak.     

Tolerance of estuarine benthic diatoms to high concentrations of ammonia,nitrite ion,nitrate ion and orthophosphate

A carbon-14 assimilation method was used to determine action spectra and photosynthesis versus irradiance (P versus I) curves of natural populations of phytoplankton and zooxanthellae from a coral reef fringing Lizard Island in the Australian Barrier Reef. The action spectra were related to the phytoplankton species composition. The curves showed shade adaptation in phytoplankton from deeper waters and in the zooxanthellae. Rates of photosynthesis of zooxanthellae were shown to be highly but variably dependent on their host organisms. Photosynthetic production by zooxanthellae was about 0.9 gC m^-2 day^-1, which is about three times higher than phytoplankton production in the waters close to the reef.     

黄海春季表层叶绿素和初级生产力及其粒径结构研究

根据2006年4月对黄海浮游植物分级叶绿素及初级生产力的调查,研究了黄海叶绿素及初级生产力的水平分布及粒级结构特征,并分析了其主要影响因素。黄海海域调查站位表层叶绿素a质量浓度变化范围为0.20～4.94μg·L-1,平均值为0.96μg·L-1。叶绿素最大值出现在临近长江口的站位。叶绿素分级结果表明黄海春季以粒径〉5μm的浮游植物占优势。黄海表层初级生产力的变化范围为2.03～15.64mg·m-3·h-1,平均值为6.08mg·m-3·h-1。其中南黄海海域初级生产力平均为6.58mg·m-3·h-1,北黄海海域初级生产力平均为4.92mg·m-3·h-1。高值区分布在南黄海中部。受水体透明度的影响,低值区出现在临近长江口的站位。断面站位分析表明浮游植物初级生产力由北向南逐步升高,温度随纬度的变化是南北海域初级生产力水平差异的主要原因。由于粒径较小（〈5μm）的浮游植物单位叶绿素具有较高的碳固定能力,调查期间整个海区初级生产力以粒径〈5μm的浮游植物贡献为主。     

The impact of grazing by microzooplankton in northern Hiroshima Bay,the Seto Inland Seam,Japan

The abundance of microzooplankton and their grazing impact on phytoplankton were studied using the dilution technique from May 1990 to November 1991 in northern Hiroshima Bay, a typical eutrophic area in the Seto Inland Sea. Microzooplankton, dominated in number by tintinnid ciliates, were abundant from June to September when chlorophyll-a concentrations were high. Maximum density of microzooplankton ranged from 3.8×10³ to 25.4×10³ ind l^-1. During the period of investigation, mean microzooplankton density and mean chlorophyll-a concentration of the <20-m fraction increased toward the inner region of the bay. The microzooplankton grazing on phytoplankton increased from summer to early autumn, and decreased from late autumn to winter. At an offshore station, the annual means of the daily grazing loss for total chlorophyll-a and the chlorophyll-a of the <20-m fraction were 12 and 15% of the initial standing stock, respectively. At an estuarine station, the microzooplankton grazed 19 and 29% of the total and <20-m initial standing stock, respectively. The quantity of grazed chlorophyll-a correlated positively and linearly with the potential production of chlorophyll-a at both stations. The quantity of chlorophyll-a grazed by microzooplankton and the potential production of chlorophyll-a were nearly equivalent in the <20-m fraction at the estuarine station. This suggests that the microzooplankton assemblage was able to consume almost all the nanoplankton newly produced in the eutrophic estuary.     

Effect of temperature and light on the growth of micro- nano- and pico-plankton: impact on algal succession

The influence of light and temperature on the phytoplankton succession in a temperate sea area was investigated in laboratory experiments with natural assemblages of micro-, nano-, and picoplankton collected from the northern Baltic Sea during 1989 and 1990. Respiration increased from 0 to 30°C in all groups of phytoplankton, while gross photosynthesis stabilised at 10 to 15°C. Light saturation occurred at 25 to 75 mol quanta m^-2 s^-1, indicating low light adaptation of the algae. Picoplankton showed the strongest temperature response, and at temperatures above 10°C picoplankton obtained a higher biomass specific photosynthesis than that of the other groups. Different light treatments had no effect on the species composition in experiments with natural algal assemblages, while different temperature conditions had a marked effect. With a temperature increase from 0 to 10°C, the algal community changed from a typical spring community, with diatoms and dinoflagellates, to a summer community, dominated by mixotrophic nanoflagellates. The small, or often non-existing, autumn bloom in the sea area studied, can be explained by short day lengths combined with relatively high temperatures, causing high community respiration rates and low gross photosynthesis, resulting in a negative energy balance. The net energy gain depends on a differential temperature effect on gross photosynthesis and endogenous respiration in various plankton groups. This gives the phytoplankton groups diverse competitive advantages during different seasons and thus may be an important factor in controlling algal succession.     

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.     

Species-specific grazing rates of heterotrophic dinoflagellates in oceanic waters,measured with a dual-label radioisotope technique

A dual-isotope method was developed to measure grazing rates and food preferences of individual species of heterotrophic dinoflagellates from natural populations, collected from the Slope, Gulf Stream, and Sargasso Sea and from a transect from Iceland to New England, in 1983. The isotope method measures the grazing rates of microzooplankton which cannot be separated in natural populations on the basis of size. Tritiated-thymidine and ¹⁴C-bicarbonate were used to label natural heterotrophic and autotrophic food, respectively. Nine oceanic dinoflagellate species in the genera Protoperidinium, Podolampas, and Diplopsalis fed on both heterotrophic and autotrophic food particles with clearance rates of 0.4 to 8.0 l cell^-1 h^-1, based on ³H incorporation, and 0.0 to 28.3 l cell^-1 h^-1, based on ¹⁴C incorporation. Two dinoflagellate species, Protoperidinium ovatum and Podolampas palmipes, fed only on ³H-labelled food particles. Several species of dinoflagellates fed on bacteria (<1 m) which had been prelabelled with ³H-thymidine. The clearance rates of heterotrophic dinoflagellates and ciliates were similar and within the range of tintinnid ciliate clearance rates reported in the literature. As heterotrophic dinoflagellates and ciliates can have comparable abundances in oceanic waters, we conclude that heterotrophic dinoflagellates may have an equally important impact as microheterotrophic grazers of phytoplankton and bacteria in oceanic waters.Partially supported by a grant from the National Science Foundation, OCE-81-17744     

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.