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.     

Sinking-rate response of natural assemblages of temperate and subtropical phytoplankton to nutrient depletion

Phytoplankton assemblages were collected during spring blooms in 1982 in Washington State and in Hawaii. Sinking rate responses of these assemblages were examined under nitrate, phosphate, and silicate depletion. Ambient nutrient concentrations, chlorophyll concentrations, photosynthetic rates, sinking rates, and floristic compositions were determined. Under nutrient-replete conditions, the temperate assemblage, composed primarily of large centric diatoms, had a sinking rate of 0.96 m d^-1; sinking rates did not change appreciably over 4 d without nitrate. Without phosphate or silicate, the sinking rates remained constant for 3 d and then increased after biomass indices began to decline. These findings illustrate the potential importance of phosphate or silicate depletion to the sedimentation of spring-bloom diatom populations. The subtropical assemblage, composed primarily of diatoms, coccolithophorids, and dinoflagellates, had an initial sinking rate of 0.22 m d^-1 and did not display substantial sinking rate changes in the absence of nitrate, phosphate or silicate. Floristic data consistently showed a proliferation of pennate diatoms, which had lower settling rates than centric diatoms. Growth and sedimentation patterns indicated a competitive advantage for pennate diatom components of subtropical assemblages; this in turn may limit phytoplankton sedimentation losses in such ecosystems.     

Distribution of phytoplankton communities in relation to the large-scale hydrographical regime in the Fram Strait

In June and July 1984 phytoplankton distribution was investigated in the Fram Strait between Greenland and Svalbard. Chlorophylla, particulate organic carbon, nitrate and phytoplankton species composition were determined from six different depths in the upper 200 m of the water column. Multivariate analysis methods were applied to identify phytoplankton communities in relation to different hydrographic regimes. Three main domains could be distinguished in terms of both hydrography and biology: (1) the East Greenland shelf polynya with a high biomass mainly produced by chain-forming diatoms, (2) the ice-covered East Greenland Current with an extremely low standing stock dominated by flagellates and (3) the marginal ice zone with a biomass maximum in 20 to 40 m depth formed by diatoms, dinoflagellates andPhaeocystis pouchetii.     

Effects of silicate depletion on photosynthesis by diatoms in the plume of the Hudson River

Silicate depletion was observed during a bloom of netplankton diatoms. Netplankton chlorophyll a increased over the same salinity range and at the same rate that silicate decreased. Silicate depletion coincided with a decrease in the apparent Si:N uptake ratio as well as a change in the magnitude and diurnal phasing of light saturated photosynthesis (P _m ^B ) by netplankton diatoms. Nanoplankton P _m ^B was unaffected by silicate depletion and increased with temperature. Consequently, nanoplankton P _m ^B eventually exceeded netplankton P _m ^B while netplankton biomass was still increasing relative to nanoplankton biomass.     

Phytoplankton sinking rates in oligotrophic waters off Hawaii,USA

The sinking rates in two size fractions of natural phytoplankton were measured over much of the photic zone in a subtropical environment. At 24, 40 and 71 m, sinking rates (± SD) of the 3 to 20 m fractions were 0.72±0.05, 0.83±0.05, and 0.34±0.04 m · d^-1, respectively, and rates for the 20 to 102 m fraction were 1.50±0.21, 1.65±0.14, and 0.95±0.22 m · d^-1, respectively. At all depths sampled, the 20 to 102 m size fraction was observed to sink significantly (P<0.01) faster than the 3 to 20 m fraction. considering vertical variability, both size fractions were observed to have significantly (P<0.01) lower sinking rates at 71 m than at more shallow depths. The finding of lower sinking rates in the region which lies just above the subsurface chlorophyll maximum provides empirical support for the hypothesis that variations in phytoplankton buoyancy may be related to the maintenance of this feature which is typical in oceanic environments. Analysis of sinking rate traces describes the distributions of specific sinking velocities for each size fraction and their variation.Oceanic Institute Contribution No. 162     

Seasonal studies on the relative importance of different size fractions of phytoplankton in Narragansett Bay (USA)

The composition and productivity of four different size-fractions (<20, 20 to 60, 60 to 100, >100 μm) of the phytoplankton of lower Narragansett Bay (USA) were followed over an annual cycle from November, 1972 to October, 1973. Diatoms dominated the population in the winter-spring bloom and in the fall, the summer population was dominated by flagellates. The nannoplankton (<20 μm) were the most important, accounting for 46.6% of the annual biomass as chlorophyll a and 50.8% of the total production. The relative importance of the different fractions showed a marked seasonality. During the winter-spring and fall blooms the netplankton fractions (>20 μm) were the most important. Nannoplankters domnated in the summer. The yearly mean assimilation numbers for the different fractions were not signfficantly different. During the winter-spring bloom, however, the assimilation numbers for the netplankters were significantly higher than those for the nannoplankton fraction. Temperature accounted for most of the variability in assimilation numbers; a marked nutrient stress was observed on only two occasions. Growth rates calculated from ¹⁴C uptake and adenosine triphosphate (ATP)-cell carbon were generally quite high; maxima were >1.90 doublings per day during blooms of a flagellate in the summer and of Skeletonema costatum in the fall. The series of short cycles observed in which the dominant species changed were related to changes in the physiological state of the population. Higher growth rates were generally observed at times of peak phytoplankton abundance while lower growth rates were observed between these peaks. The high growth rates and assimilation numbers usually found suggest that the phytoplankton in lower Narragansett Bay was not generally nutrient-limited between November, 1972 and October, 1973. Nutrient regeneration in this shallow estuary, therefore, must be very rapid when in situ nutrient levels are low.     

Light-saturated photosynthesis by phytoplankton size fractions in the New York bight,USA

The relationships between netplankton and nanoplankton assimilation numbers, temperature, and major nutrient concentrations were studied and evaluated in the context of seasonal patterns in the biomass of these phytoplankton size fractions. Netplankton and nanoplankton blooms typically occur during late winter (2° to 8°C) and summer (18° to 24°C), respectively. Variations in nanoplankton and netplankton assimilation numbers were not statistically related to the development or collapse of specific blooms based on weekly sampling, but assimilation numbers were higher during the bloom periods than during transition periods of rapid temperature change (8° to 18°C). Differences in the assimilation numbers between size fractions could account for the dominance of the nanoplankton fraction during the summer bloom period but not for the dominance of netplankton during the winter bloom period. Nanoplankton and netplankton assimilation numbers were exponential functions of temperature between 8° and 24°C and 8° and 20°C, respectively. Below 8°C the assimilation numbers of both fractions were higher than expected on the basis of temperature. Above 20°C netplankton assimilation numbers declined with temperature. Netplankton and nanoplankton assimilation numbers were occasionally correlated with dissolved inorganic nitrogen concentrations from less than 1.0 to more than 15 g-at l^-1. Under these conditions, nanoplankton growth rates (calculated from assimilation number and carbon:chlorophyll) were higher and increased more rapidly with dissolved inorganic nitrogen than netplankton growth rates.     

Parameters of light-dependent photosynthesis for phytoplankton size fractions in temperate estuarine and coastal environments

Photosynthetic parameters for netplankton (>22 m) and nanoplankton (<22 m) varied over similar ranges but exhibited different seasonal and geographic patterns of variation. Nanoplankton a was relatively constant (0.06 mg C [mg Chl · h]^-1 [E m^-2 s^-1]^-1), but P _m ^B (mg C [mg Chl · d]^-1) was an exponential function of temperature independent of nutrient concentration and vertical stability in the euphotic zone. The temperature function gives a P _m ^B of 24 at 25°C for nanoplankton growing in an estuarine environment characterized by high nutrient concentrations and a shallow, stratified euphotic zone. Variations in netplankton a and P _m ^B were less predictable and were not correlated with temperature, nutrients or vertical stability. Chain forming diatoms with small cells were able to achieve high (0.10 to 0.15) and P _m ^B (20 to 24) that were 3 to 5 times higher than large-celled diatoms and dinoflagellates were able to achieve.     

Dynamique du phytoplancton et matière organique particulaire dans la zone euphotique de l'Atlantique Equatorial

At two fixed stations in the Equatorial Atlantic Ocean (0°–4° W), the physical, chemical and biological properties of the euphotic layer were determined for 14 d (Station A: 5–18 February, 1979) and 13 d (Station B: 20 October–7 November, 1979), respectively. The stability of the water column allowed comparison of 3 different “systems”: (i) a well-illuminated and nitrate-depleted mixed layer; (ii) a chlorophyll maximum layer (chl a _max) in the thermocline which is poorly illuminated (6.3% of surface irradiance); (iii) a well-illuminated but nitrate-rich (>0.9 μg-at l^-1) mixed layer. In each layer the particulate organic carbon (COP), nitrogen (NOP) and phosphorus (POP) contents were measured and compared with the phytoplankton biomass. In the chlorophyll maximum layer, the phytoplankton biomass contributed significantly to the total particulate organic matter (between 55 and 75%). In the nitrate-depleted mixed layer, the results varied according to whether the regression technique [COP=f(chl a)] was used, or the chl a synthesis during the incubation of the samples. With the former technique, the phytoplankton carbon (C _p) content appeared minimal, because the y intercept, computed using all the data of the water column, was probably overestimated for this layer. POP would be more associated with living protoplasm than with carbon and nitrogen in the three layers. In the chlorophyll a maximum layer it constitutes a valuable detritus-free biomass measurement, since 80% of the POP consist of phytoplankton phosphorus. The assimilation numbers (NA=μg C μg chl a ^-1 h^-1) were high in all three layers, but the highest values were recorded in the nitrate-depleted mixed layer (NA=15 μg C μg chl a ^-1 h^-1). In the chlorophyll maximum layer, light would be a limiting factor during incubation: between 10²⁵ and 8.10²⁴ quanta m^-2 d^-1 NA and light are positively correlated independant of nitrate concentration. The growth rates of phytoplankton (μ) were estimated and compared to the maximum expected growth rate. Our main conclusion was that despite very low biomass and nutrient content, the mixed layer was in a highly dynamic state, as evidenced by high rates of phytoplankton growth and short nutrient turnover times (1 d or less for PO-P₄ in the mixed layer versus 3 d in the thermocline). The presence of nitrate in the water column allows the development of a higher phytoplankton biomass but does not increase growth rate.     

Spring phytoplankton in the Subarctic North Pacific Ocean

The report includes quantitative and qualitative data on the phytoplankton from the First Canadian Transpacific Oceanographic Cruise from March to May 1969. Nanoplanktonic species (<20 m) were numerically dominant and, together with nertic diatoms, made up the larger portion of the biomass at all stations outside the Western Subarctic and Alaskan Gyres, where Denticulopsis seminae predominated. The nanoplankton consisted mainly of haptophycean, dinophycean and cryptophycean flagellates. Tropical oceanic dinoflagellates and diatoms were abundant in the net samples from Station 17 in the warm Kuroshio current but absent from Stations 24–27 in the Western Subarctic Gyre, Stations 2, 37–39 in the Alaskan Gyre, in which cold water oceanic species predominated, and Stations 31 and 33, south of the Aleutians. Subsurface maxima were common on the outward bound (westerly) leg in March.     

Species-specific sedimentation and sinking velocities of diatoms

Sedimentation rates were determined for various diatom species, and both average and maximum sinking velocities of sedimenting diatoms were calculated during a spring bloom investigation in the central Baltic Sea in 1986. Up to 25 and 50% of theChaetoceros spp. andThalassiosira levanderi populations, respectively, sedimented daily. Daily sedimentation rates of other diatoms, dinoflagellates andMesodinium rubrum, however, were less than 1% of their respective standing stocks. TheT. levanderi population was divided into two subpopulations: while one was sinking, the second was actively dividing (recognizable by paired-cell stages) with a specific growth rate of >0.2 to 0.3 d^–1. These paired cells were never found in sediment trap samples. The average sinking velocity ofChaetoceros spp. was 15 to 30 m d^–1; that ofT. levanderi was higher. The maximum sinking velocity of cells was at least 70 m d^–1. According to these observations, the formation of aggregates (which enhances sinking velocity), and their sedimentation, represent a highly selective process. This indicates that diatom aggregates do not act as roving filters, sweeping the water clear while sinking.     

Particulate protein-nitrogen in North Atlantic surface waters

Depth profiles of particulate protein-nitrogen at 4 oceanic and 2 upwelling stations in the North Atlantic Ocean were measured by a new fluorometric method. The protein-nitrogen in the upper 20 m ranged from 0.19 to 1.61 μg-at N/1 at the oceanic stations and from 0.43 to 3.54 μg-at/1 at the upwelling stations. The mean values in the euphotic zone were 0.54 μg-at N/1 for the oceanic stations and 1.70 μg-at N/1 for the upwelling stations. The ratio of protein-nitrogen to chlorophyll at the two sets of stations was 2.83 and 0.54 μg-at N/μg chlorophyll, respectively. Regression analysis of the pooled data yielded a detritus and zooplankton-free ratio of 0.38 μg-at N:μg chlorophyll. Calculations of the phytoplankton protein-nitrogen, based on this ratio, suggest that in the oceanic water only 20% of the sestonic protein-nitrogen is associated with the phytoplankton. In the upwelling waters, the phytoplankton may account for 65% of the sestonic proteinnitrogen.     

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.     

Production of picoplankton and small nanoplankton in the Celtic Sea

A significant proportion of the total primary production in the Celtic Sea (50°30′N; 07°00′W) has been found to be due to picoplankton and small nanoplankton. In July, August and October, 1982, 20 to 25% of the ¹⁴C fixed in primary production was in organisms >5 μm, 35 to 40% was in organisms <5–1 μm and 20 to 30% was in organisms<1 μm. Bacterial production was estimated by the incorporation of ³H and would account for less than 10% of the production in the <1–>0.2 μm fraction; therefore, production in the <1–>0.2 μm fraction was the result of photosynthesis per se by picoplankton and could not have been due to heterotrophic bacteria utilizing exudates from larger phytoplankton. Time-course experiments demonstrated some transfer of label from the <1–>0.2μm fraction to the >5 μm fraction, presumably by grazing, but again most of the production in this fraction was the result of photosynthesis by organisms larger than 5 μm and was not due to grazing by heterotrophic microflagellates on smaller phytoplankton.     

Vertical fluxes and ecological significance of organic materials during the phytoplankton bloom during austral summer in Breid Bay,Antarctica

A mooring system consisting of an in situ fluorometer at a depth of 50 m and a time-series sediment trap at a depth of 110 m was deployed at the sediment trap site (70°11.536'S; 24°18.679'E; water depth: 300 m) in Breid Bay, Antarctica in austral summer from 28 December 1985 to 13 February 1986. Sinking particles, consisting of diatoms (mainly Thalassiosira antarctica), were analyzed for organic materials, stable carbon and nitrogen isotopes. Vertical fluxes of organic carbon and nitrogen were determined to be within the ranges of 12.3 to 116 mg C m^-2 d^-1 and 1.79 to 15.4 mg N m^-2 d^-1, respectively, with maxima in the middle of January 1986, after which time the organic carbon and nitrogen fluxes tended to decrease with a steep gradient. High values of ¹³C were found in the organic matter of the sinking particles collected before the middle of January, indicating that the organic matter was derived from the diatoms in the logarithmic phase of their growth. Increased abundance of glucose was found in the water-extractable carbohydrate, which was one of the sinking particles collected in the middle of January. This fact clearly indicated that the diatoms were no longer in the growth phase but rather in the stationary phase, because reserved glucan as well as various cellular organic materials were reportedly accumulated within the algal cells in the stationary phase. Fifteen species of protein amino acids with trace amounts of -alanine, -aminobutyric acid and ornithine were found in the sinking particles upon acid hydrolysis, but the amino acid composition of these samples had not been affected much by biological agents. The essential amino acid index was calculated for the sinking particles collected in the course of the sediment trap experiment. The indices obtained indicated that the sinking particles collected in Breid Bay were more ecologically significant than the sinking and suspended particles from deep waters.     

Significance of nanoplankton in the Chesapeake Bay estuary and problems associated with the measurement of nanoplankton productivity

Over a 2-year program of monthly cruises covering the entire Chesapeake Bay (USA), the phytoplankters which passed 35 μm mesh were responsible for 89.6% of the phytoplankton productivity. On a single summer cruise, the <35 μm phytoplankton fraction was responsible for 93.4% of the chlorophyll a and 100% of the primary productivity. The <10 μm fraction was responsible for 81.3% of the chlorophyll a and 94% of the productivity. The difference in biomass in the <35 μm and the <10 μm fractions was significant (P=0.025), but no significant difference in the productivity could be demonstrated. Laboratory experiments demonstrated that recently assimilated carbon can be lost with gravity screening. Considering both this and the effect of herbivorous zooplankters enclosed in productivity incubations, a prescreening rather than postscreening technique is recommended for studying nanoplankton productivity.     

The spring phytoplankton bloom in Lindåspollene,a land-locked Norwegian fjord. II. Biomass and activity of net- and nanoplankton

From February 24 to April 24, weekly samples were collected at fixed depths at one station in Lindåspollene, a land-locked Norwegian fjord. Adenosine triphosphate (ATP), chlorophyll a, phaeophytin, ¹⁴C assimilation, and respiratory activity [electron transport system (ETS) activity] were measured in the net- (>30 m) and nanoplankton. Netplankton contained on the average 48% of the total chlorophyll a and 56% of the ATP, but contributed only 7% to the total carbon assimilation and 11% to the ETS activity. The assimilation numbers for net- and nanoplankton ranged from 0 to 1.2 and from 1.5 to 13.2, respectively. At the oxygen/hydrogen sulphide interface, high concentrations of ATP, but not of chlorophyll a, were found in the nanoplankton fraction. Netplankton algae grew actively only in the first phase of the bloom, and nanoplankton predominated later, apparently due to low nutrient concentrations. During the bloom, Skeletonema costatum made up the main part of the biomass. The number of cells in the chains decreased throughout the bloom, possibly reflecting the lowered silicate content. It appeared that only nanoplankton were grazed by zooplankton, while netplankton sank to the bottom and represented input to the benthos.     

Diurnal rhythms in netplankton and nannoplankton assimilation ratios

Surface nannoplankton and netplankton photosynthetic rates and chlorophyll a concentrations were measured 3 h before and 3 h after local apparent noon at 36 stations in the eastern Pacific. Morning and afternoon nannoplankton and netplankton photosynthetic capacities were plotted against chlorophyll a concentrations. Comparison of am (morning) and pm (afternoon) regression coefficients revealed that neither netplankton nor nannoplankton photosynthetic efficiencies varied diurnally in temperate California Current waters. However, in oligotrophic tropical surface waters where the photosynthetic rates of both fractions were nitrogen limited, netplankton assimilation ratios exhibited an afternoon maximum, while nannoplankton efficiencies exhibited a morning maximum. The netplankton followed the same pattern in eutrophic tropical surface waters with high nitrate concentrations, but nannoplankton assimilation ratios were highest in the afternoon. Assuming that midday light intensities inhibit surface photosynthesis, it was concluded that assimilation ratios will reach a mid-morning or mid-afternoon maximum depending upon the relative importance of netplankton and nannoplankton productivity and the degree to which nutrients are limiting.     

A detailed analysis of carbohydrates in marine particulate matter

Detailed profiles of organic carbon, organic nitrogen, carbohydrate and proteinous amino acid have been determined in particulate matter from various depths at a station in Sagami Nada off eastern Honshu, Japan. The profiles suggest that carbohydrate is decayed from particulate matter more rapidly than proteinous amino acid in the euphotic zone, while the latter is removed to a higher extent than the former in deep waters,which results in the increase of the C/N values of particulate matter in the depths. All of the particulate matter gave D-galactose, D-glucose, D-mannose, D-xylose and D-glucuronic acid upon acid hydrolysis. Vertical change of the monosaccharide composition of this particulate matter indicates that only D-glucose and its polymers are preferentially removed from it during its descent. Carbohydrates from the particulate matter of 20 m depth were fractionated into water-soluble and insoluble carbohydrates. Detailed analysis of these fractions indicates that 1,3-glucan and its related low molecular weight carbohydrates from the water-soluble fraction decay between 50 and 300 m depth, to leave waterinsoluble carbohydrates, immune to biological attack during the course of further sinking. On the basis of these facts, the effects of the biochemical nature of these carbohydrates on the vertical change of the particulate carbohydrate distribution are discussed.     

Seasonal study of phytoplankton pigments and species at a coastal station off Sydney: Importance of diatoms and the nanoplankton

Phytoplankton pigments and species were studied at a coastal station off Sydney (New South Wales, Australia) over one annual cycle. Sudden increases in chlorophyll a (up to 280 mg m^-2), due to short-lived diatom blooms, were found in May, July, September, January and February. These were superimposed upon background levels of chlorophyll a (20 to 50 mg m^-2), due mostly to nanoplankton flagellates, which occurred throughout the year. The nanoplankton (<15 m) accounted for 50 to 80% of the total phytoplankton chlorophyll, except when the diatom peaks occurred (10 to 20%). The annual cycle of populations of 16 dominant species-groups was followed. Possible explanations as to alternation of diatom-dominated and nanoplankton-dominated floras are discussed. Thin-layer chromatography of phytoplankton pigments was used to determine the distribution of algal types, grazing activity, and phytoplankton senescence in the water column. Chlorophyll c and fucoxanthin (diatoms and coccolithophorids) and chlorophyll b (green flagellates) were the major accessory pigments throughout the year, with peridinin (photosynthetic dinoflagellates) being less important. Grazing activity by salps and copepods was apparent from the abundance of the chlorophyll degradation products pheophytin a (20 to 45% of the total chlorophyll a) and pheophorbide a (10 to 30%). Chlorophyllide a (20 to 45%) was associated with blooms of Skeletonema costatum and Chaetoceros spp. Small amounts of other unidentified chlorophyll a derivatives (5 to 20%) were frequently observed.