Seasonal and interannual variability of size-fractionated phytoplankton biomass and community structure at station Kerfix, off the Kerguelen Islands, Antarctica

Time series of phytoplankton biomass and taxonomic compositionhave been obtained for the 3 years 1992, 1993 and 1994 in thenorthern part of the Southern Ocean (station Kerfix, 5040'S,6825;E) Autotrophic biomass was low throughout the year (<0.2mg m^–3 except during a short period in summer when a maximumof 1.2 mg chlorophyll (Chl) a m– was reached. During winter,the integrated biomass was low (<10 mg m^–2) and associatedwith deeply mixed water, whereas the high summer biomass (>20mg m^–2) was associated with increased water column stability.During summer blooms, the >10 µ;m size fraction contributed60% to total integrated biomass. Large autotrophic dinoflagellates,mainly Prorocentrum spp., were associated with the summer phytoplankton maxima and accounted for >80% of the total autotrophcarbon biomass. In November and December, the presence of thelarge heterotrophic dinoflagellates Protoperidinium spp. andGyro dinium spp. contributed a high proportion of total carbonbiomass. During winter, the <10 µm size fraction contributed80% of total Chi a biomass with domination of the picoplanktonsize fraction. The natural assemblage included mainly nakedflagellates such as species of the Prasinophyceae, Cryptophyceaeand Prymnesiophyceae. During spring, picocyanobacteria occurredin sub-surface water with a maximum abundance in September of106 cells 1^–1     

Community structure, biomass and productivity of size-fractionated summer phytoplankton populations in lower Narragansett Bay, Rhode Island

Seventeen size-fractionation experiments were carried out duringthe summer of 1979 to compare biomass and productivity in the< 10, <8 and <5 µm size fractions with that ofthe total phytoplankton community in surface waters of NarragansettBay. Flagellates and non-motile ultra-plankton passing 8 µmpolycarbonate filters dominated early summer phytoplankton populations,while diatoms and dinoflagellates retained by 10 µm nylonnetting dominated during the late summer. A significant numberof small diatoms and dinoflagellates were found in the 10–8µm size fraction. The > 10 µm size fraction accountedfor 50% of the chlorophyll a standing crop and 38% of surfaceproduction. The <8 µm fraction accounted for 39 and18% of the surface biomass and production. Production by the< 8 µm fraction exceeded half of the total communityproduction only during a mid-summer bloom of microflagellates.Mean assimilation numbers and calculated carbon doubling ratesin the <8 µm (2.8 g C g Chl a^–1 h^–1; 0.9day^–1)and<5 µm(1.7 g C g Chl a^–1h^–1; 0.5day^–1)size fractions were consistently lower than those of the totalpopulation (4.8 g C g Chl a^–1 h^–1; 1.3 day^–1)and the <10 µm size fraction (5.8 g C g Chl a^–1h^–1; 1.4 day ^–1). The results indicate that smalldiatoms and dinoflagellates in fractionated phytoplankton populationscan influence productivity out of proportion to their numbersor biomass. ¹Present address: Australian Institute of Marine Science, P.M.B.No. 3, Townsville M.S.O., Qld. 4810, Australia.     

Seasonal photosynthetic activity of autotrophic picoplankton in Lake Kinneret, Israel

Autotrophic picoplankton populations in Lake Kinneret are composedof picocyanobacteria and picoeukaryotes. Overall, the ratesof photosynthetic carbon fixed by autotrophic picoplankton duringthis study were low (0.01–1.5 mg Cm^–3 h^–1).The highest chlorophyll photosynthetic activity of the <3µm cell-size fraction was found in spring, when picoeukaryotespredominated and in addition small nanoplankton passed throughthe filters. The maximum cell-specific photosynthetic rate ofcarbon fixation by picocyanobacteria and picoeukaryotes was2.5 and 63 fg C cell^–1 h^–1, respectively. The highestspecific carbon fixation rate of autotrophic picoplankton was11 µg C µg^–1 Chl h^–1 The proportionalcontribution of autotrophic picoplankton to total photosynthesisusually increased with depth. Picocyanobacteria collected fromthe dark, anaerobic hypolimnion were viable and capable of activephotosynthesis when incubated at water depths within the euphoticzone. Maximum rates of photosynthesis (P_max) for picocyanobacteriaranged from 5.4 to 31.4 fg C cell^–1 h^–1 with thehighest values in hypolimnetic samples exposed to irradiance.Photosynthetic efficiency (     

Ammonium excretion and glutamate dehydrogenase activity of zooplankton in Great South Bay, New York

In Great South Bay, nanoplankton, (<20 sµm) accountedfor the largest fraction (56%) of zooplankton glutamate dehydrogenase(GDH) activity over a one year period. Microzooplankton (20–200µm) and macrozooplankton (>200 µm) accountedfor 20% and 24%, respectively. Total zooplankton ammonium regenerationin Great South Bay could account for 74% of the ammonium requirementby phytoplankton in winter, but in summer when phytoplanktondemand was greater, and zooplankton population was low, it suppliedless than 5%. This study suggests that the smallest zooplanktonfraction, less than 20 µm, can be the most important asregards nitrogen regeneration in estuarine environments. MacrozooplanktonGDH activity in Great South Bay ranged from 0.18 mg atoms NH⁺₄-Nm^–3 d^–1 in winter to 3.34 mg atoms NH⁺₄-N m^–3d^–1 in spring. Over an annual period, the averaged GDH/excretionratio was 20.4 3.5 (n = 10), and this ratio agrees well withobservations by other investigators. Observed macrozooplanktonexcretion rates showed a strong correlation with the excretionrates indirectly estimated from GDH activities. The GDH/excretionratio seems to vary depending on the internal physiologicalstates of zooplankton as well as food availability.     

Size and species-specific primary productivity and community structure of phytoplankton in Tokyo Bay

Combined methods of size fractionation and single-cell isolationwere used to investigate the seasonal variation of phytoplanktondynamics in Tokyo Bay with an emphasis on primary productivity.Red tides occurred in Tokyo Bay from spring to autumn; a diatom,Skeletonema costatum, and a raphidophycean, Heterosigma akashiwo,were the most important primary producers. Small diatoms andflagellates, including these species, were dominant and showedrapid changes of phytoplankton community structure within severaldays in summer. The nanoplankton (3–20 µm) fractioncontributed most to chlorophyll a concentration and primaryproductivity during spring to autumn, whereas the microplankton(>20 µm) contribution was remarkable in winter. Picoplankton(<3 µm phytoplankton) remained relatively constantthroughout the year. A significant reverse relationship wasobtained between assimilation rate and chlorophyll a contentfor the total and nanoplankton population; the assimilationrate was high at the initial phase of the bloom, then decreasedto a minimum level at the peak of the bloom. Factors controllingthe reduction of assimilation rates at the peak, and changesin phytoplankton community structure, are discussed.     

Seasonal patterns of urea regeneration by size-fractionated microheterotrophs in well-mixed temperate coastal waters

Urea regeneration by size-fractionated plankton was measuredover an annual cycle at a coastal station in the permanentlywell-mixed waters of the western English Channel. Rates of urearegeneration in the <200 µm fraction varied from 0.6to 20.6 nmol N L^–1 h^–1. Regeneration rates werelowest in winter and highest in summer. The ratio of the ratesof regeneration to uptake of urea was close to 1 on all time(seasonal and nycthemeral), and space (vertical) scales indicatingthat regeneration by microheterotrophs supplied the totalityof urea used by phytoplankton. On an annual basis, urea regeneratedby the microheterotrophs (0.98 mol N m^–2 year^–1)was equivalent to 33% of the total regenerated N (urea + ammonium).The major part of urea regeneration was due to the nanoplankton(51%) and microplankton fractions (36%). Regeneration of ureain the picoplankton was detectable only from April to Octoberand represented, on an average, 25% of the total urea regeneratedduring this period. Urea regeneration in micro- and nanoplanktonfractions was mainly associated with ciliates and in the picoplanctonfraction with bacteria.     

Seasonal variations of size-fractionated phytoplankton along the salinity gradient in the York River estuary, Virginia (USA)

The dynamics of phytoplankton size structure were investigatedin the freshwater, transitional and estuarine zones of the YorkRiver over an annual cycle. The contribution of large cells(microplankton, >20 µm) to total concentrations ofchlorophyll a increased downstream during winter, whereas thatof small cells (nanoplankton, 3–20 µm; picoplankton,<3 µm) increased downstream during summer. In the freshwaterregion, the contribution of micro phytoplankton to total concentrationsof chlorophyll a was significant during warm seasons (springand summer) but not during colder seasons (winter), whereasthe contribution of small-sized cells (especially picoplankton)increased during cold seasons. Temperature, light and high flushingrate appear to control phytoplankton community structure inthe freshwater region. In the transitional region, nano-sizedcells dominated the phytoplankton population throughout allseasons except during the spring bloom (April) when the chlorophylla concentration of micro phytoplankton increased. Size structurein the transitional region is most likely regulated by lightavailability. In the mesohaline region, nano- and pico-sizedcells dominated the phytoplankton population during the summerbloom, whereas micro-sized cells dominated during the winterbloom. Factors controlling phytoplankton community size structurein the mesohaline zone may be riverine nitrogen input, temperatureand/or advective transport from up-river. Based on these results,the spatial and seasonal variations in size structure of phytoplanktonobserved on the estuarine scale may be determined both by thedifferent preferences for nutrients and by different light requirementsof micro-, nano- and picoplankton. The results suggest thatanalyses of phytoplankton size structure are necessary to betterunderstand controls on phytoplankton dynamics and to bettermanage water quality in river-dominated, estuarine systems.     

Carbon dynamics in the 'grazing food chain' of a subtropical lake

Studies were conducted over a 13 month period at four pelagicsites in eutrophic Lake Okeechobee, Florida (USA), in orderto quantify carbon (C) uptake rates by size-fractionated phytoplankton,and subsequent transfers of C to zooplankton. This was accomplishedusing laboratory ¹⁴C tracer methods and natural plankton assemblages.The annual biomass of picoplankton (<2 µm), nanoplankton(2–20 µm) and microplankton (<20 µm averaged60, 389 and 100 µg C 1^–1 respectively, while correspondingrates of C uptake averaged 7, 51 and 13 µg C1^–1h^–1. The biomass of microzooplankton (40–200 µm)and macrozooplankton (<200 µm averaged 18 and 60 µgC 1^–1, respectively, while C uptake rates by these herbivoregroups averaged 2 and 3 µg C 1^–1 h^–1. Therewere no strong seasonal patterns in any of the plankton metrics.The ratio of zooplankton to phytoplankton C uptake averaged7% over the course of the study. This low value is typical ofthat observed in eutrophic temperate lakes with small zooplanktonand large inedible phytoplankton, and indicates ineffectiveC transfer in the grazing food chain. On a single occasion,there was a high density (<40 1^–1) of Daphnia lumholrzii,a large-bodied exotic cladoceran. At that time, zooplanktoncommunity C uptake was <20 µg C 1^–1 h^–1and the ratio of zooplankton to phytoplankton C uptake was near30%. If D.lumholrzii proliferates in Lake Okeechobee and theother Florida lakes where it has recently been observed, itmay substantially alter planktonic C dynamics.     

Size structures of microplankton biomass and production in Jiaozhou Bay, China

Seasonal investigations of size-fractionated biomass and productionwere carried out from February 1992 to May 1993 in JiaozhouBay, China. Microplankton assemblages were separated into threefractions: pico- (0.7–2 µm), nano- (2–20 µm)and netplankton (20–200 µm). The biomass was measuredas chlorophyll a (Chi a), paniculate organic carbon (POC) andparticipate organic nitrogen (PON). The production was determinedby ¹⁴C and ¹⁵N tracer techniques. The seasonal patterns in biomass,though variable, were characterized by higher values in springand lower values in autumn and summer (for Chi a only). Theseasonal patterns in production, on the other hand, were moreclear with higher values occurring in summer and spring, andlower values occurring in autumn and winter. Averaged over thewhole study period, the respective proportions of total biomassaccounted for by net-, nano- and picoplankton were 26, 45 and29% for Chi a, 32, 33 and 35% for POC, and 26, 32 and 42% forPON. The contributions to total primary production by net-,nano- and picoplankton were 31, 35 and 34%, respectively. Therespective proportions of total NH₄⁺–N uptake accountedfor by net-, nano- and picoplankton were 28, 33 and 39% in thedaytime, and 10, 29 and 61% at night. The respective contributionsto total NO₃^–-N uptake by net-, nano- and picoplanktonwere 37, 40 and 23% in the daytime, and 13, 23 and 64% at night.Some comprehensive ratios, including C/N biomass ratio, Chla/C ratio, C uptake/Chl a ratio, C:N uptake ratio and the f-ratio,were also calculated size separately, and their biological andecological meanings are discussed.     

Diurnal and seasonal variation in the contributions of autotrophic pico-, nano- and microplankton to the primary production of an upland lake

An investigation of the diurnal variation in productivity andcontribution to production of populations of autotrophic picoplankton(0.2–2.0 µm), nanoplankton (>2 <20 µm)and microplankton (>20 µm) was carried out at monthlyintervals, from May to October 1989, in Llyn Padarn a mesotrophicupland lake in North Wales. Maximum rates and contributionsto production of the lake by autotrophic picoplankton occurredduring mid-late summer, with the highest average daily contributionfrom picoplankton (64%) recorded in September at 4 m depth.Diurnal variation in contributions from picoplankton was pronounced,with greatest input, recorded at the end of the day, duringthe period of picoplankton dominance in mid-late summer. Maximumcontribution from picoplantkon (86% of total, 9.2 mg C m^–3h^–1) was recorded in September. Nanoplankton primary productionwas of greatest significance in June and July, although levelswere lower than for picoplankton in subsequent months. Contributionsvia nanoplankton increased with depth in the lake at this time,reaching a maximum of 78% of the total at the end of the dayat 9 m depth in early July. At this time, diurnal variationin contributions via nanoplankton was considerable, with maximumphotosynthesis generally at the end of the photoperiod at depthsof 4 and 9 m. Microplankton made the greatest impact on primaryproduction during the mixed water conditions of spring and autumn,and at these times did variation in production was less thanthose of both pico and nanoplankton during summer thermal stratification.Photosynthetic capacity was lower for picoplankton than fornanoplankton and microplankton; the highest values were 5, 33and 51 mg C (mg chl a)^–1) h^–1) for pico-, nano-and microplankton, respectively. The photosynthetic efficiencyof all three size categories of phytoplankton increased withdepth. Maximum values were similar for all phytoplankton groups,between 75 and 131 mg C (mg chl a)^–1) E^–1) m² butmean levels of photosynthetic efficiency for the 6 months werelower for picoplankton than for nano- or microplankton. Ratesof carbon fixation per cell for picoplankton spanned three ordersof magnitude, varied considerably diurnally and reached maximumvalues of 484 fg C(cell)^–1) h^–1) in the afternoonin near-surface waters in the early stages of exponential populationgrowth in July. During the population maximum of picoplanktonin August and September, maximum daily values of carbon fixationper cell, assimilation number and photosynthetic efficiencywere all recorded at the end of the day. The seasonal and diurnalpatterns of production of the three size categories of planktonicalgae in Llyn Padarn were distinct. During spring, microplankton(mainly diatoms) were the dominant primary producers. As thermalstratification developed, nanoplankton were the major contributorsto phytoplanktonic production, particularly in the deeper regionsof the euphotic zone. Picoplankton made the greatest contributionto production in August and September, exhibiting maximum inputtowards the end of the light cycle. Diatoms became the majorphotosynthetic plankton in the mixed water conditions prevalentin Uyn Padarn in October.     

Productivity of picoplankton compared with that of larger phytoplankton in the subarctic region

We determined the productivity (µg C µg^–1Chi a h^–1) of size-fractionated phytoplankton in the northernNorth Pacific and the Bering Sea in summer and winter. Picoplankton(<2 µm) were more productive than larger sized phytoplankton(2–10 and 10–200 µm) in the subtropical region,where the in situ temperature was >10°C; whereas picoplanktonin the subarctic region were similar in productivity or lessproductive than larger sized plankton, where the in situ temperaturewas <10°C. The result from the subtropical region inthis study agrees with previous results from tropical and subtropical waters, which indicate that phytoplankton productivitytends to decrease with increasing cell size. The result fromthe subarctic region, however, differs from previous results.We observed a positive linear regression for in situ temperatureand picoplankton productivity, but this trend was not seen inthe larger sized phytoplankton. The results show that the productivityof picoplankton is markedly influenced by in situ temperaturecompared with that of larger sized plankton. Low tem peratureappears to account largely for the observation that the productivityof picoplankton is not significantly higher than that of largersized phytoplankton in the subarctic region.     

Comparison of east and west chlorophyll a standing stock and oceanic habitat along the Transition Domain of the North Pacific

Chlorophyll (Chl) a was measured every 10 m from 0 to 150 min the Transition Domain (TD), located between 37 and 45°N,and from 160°E to 160°W, in May and June (Leg 1) andin June and July (Leg 2), 1993–96. Total Chl a standingstocks integrated from 0 to 150 m were mostly within the rangeof 20 and 50 mg m^–2. High standing stocks (>50 mg m^–2)were generally observed westof 180°, with the exceptionof the sporadic high values at the easternmost station. Thetotal Chl a standing stock tended to be higher in the westernTD (160°E–172°30'E) than in the central (175°E–175°W)and eastern (170°W–160°W) TD on Leg 1, but thesame result was not observed on Leg 2. It was likely that largephytoplankton (2–10 and >10 µm fractions) contributedto the high total Chl a standing stock. We suggest that thehigh total Chl a standing stock on Leg 1, in late spring andearly summer, reflects the contribution of the spring bloomin the subarctic region of the northwestern Pacific Ocean. Thedistribution of total Chl a standing stock on Leg 2 was scarcelyaffected by the spring phytoplankton bloom, suggesting thattotal Chl a standing stock is basically nearly uniform in theTD in spring and summer. Moreover, year-to-year variation inthe total Chl a standing stock was observed in the western TDon Leg 1, suggesting that phytoplankton productivity and/orthe timing of the main period of the bloom exhibits interannualvariations.     

The size distribution of phytoplankton and participate organic matter in the Equatorial Atlantic Ocean: importance of ultraseston and consequences

The size fractionation of paniculate matter (<200, <35,<3 and <1 µm) has been measured in the EquatorialAtlantic Ocean at different stations. Chlorophyll a, phaeophytin,particulate carbon, nitrogen and phosphorus have been analysed.Primary production by ¹⁴CO₂ uptake was also measured with prescreeningtechnique. It appears from this study, that the pariculate matter has avery small size: 40–60% of the chlorophyll passed through1 µm Nucleopore filter, and 75–90% of the paniculatecarbon and nitrogen passed through 3 µm Nucleopore filterin offshore waters. From the atomic ratio C/N, C/P and C/chla, and primary productionvalues, the <3 µm fraction would be mainly constitutedby inactive photosynthetic organisms or partides of detritus.The 3–35 µm fraction, in contrast, would be principallyactive phytoplankton.     

Response of phytoplankton and bacteria to nutrients and zooplankton: a mesocosm experiment

Although both nutrient inputs and zooplankton grazing are importantto phytoplankton and bacteria in lakes, controversy surroundsthe relative importance of grazing pressure for these two groupsof organisms. For phytoplankton, the controversy revolves aroundwhether zooplankton grazers, especially large cladocerans likeDaphnia, can effectively reduce phytoplankton populations regardlessof nutrient conditions. For bacteria, little is known aboutthe balance between possible direct and indirect effects ofboth nutrients and zooplankton grazing. However, there is evidencethat bacteria may affect phytoplankton responses to nutrientsor zooplankton grazing through direct or apparent competition.We performed a mesocosm experiment to evaluate the relativeimportance of the effects of nutrients and zooplankton grazingfor phytoplankton and bacteria, and to determine whether bacteriamediate phytoplankton responses to these factors. The factorialdesign crossed two zooplankton treatments (unsieved and sieved)with four nutrient treatments (0, 0.5, 1.0 and 2.0 µgphosphorus (P) l^–1 day^–1 together with nitrogen(N) at a N:P ratio of 20:1 by weight). Weekly sieving with 300µm mesh reduced the average size of crustacean zooplanktonin the mesocosms, decreased the numbers and biomass of Daphnia,and increased the biomass of adult copepods. Nutrient enrichmentcaused significant increases in phytoplankton chlorophyll a(4–5x), bacterial abundance and production (1.3x and 1.6x,respectively), Daphnia (3x) and total zooplankton biomass (2x).Although both total phytoplankton chlorophyll a and chlorophylla in the <35 µm size fraction were significantly lowerin unsieved mesocosms than in sieved mesocosms, sieving hadno significant effect on bacterial abundance or production.There was no statistical interaction between nutrient and zooplanktontreatments for total phytoplankton biomass or bacterial abundance,although there were marginally significant interactions forphytoplankton biomass <35 µm and bacterial production.Our results do not support the hypothesis that large cladoceransbecome less effective grazers with enrichment; rather, the differencebetween phytoplankton biomass in sieved versus unsieved zooplanktontreatments increased across the gradient of nutrient additions.Furthermore, there was no evidence that bacteria buffered phytoplanktonresponses to enrichment by either sequestering P or affectingthe growth of zooplankton.     

Dynamics of microplankton communities at the ice-edge zone of the Lazarev Sea during a summer drogue study

Microzooplankton grazing and community structure were investigatedin the austral summer of 1995 during a Southern Ocean Drogueand Ocean Flux Study (SODOFS) at the ice-edge zone of the LazarevSea. Grazing was estimated at the surface chlorophyll maximum(5–10 m) by employing the sequential dilution technique.Chlorophyll a concentrations were dominated by chainformingmicrophytoplankton (>20 µm) of the genera Chaetocerosand Nitzschia. Microzooplankton were numerically dominated byaloricate ciliates and dinoflagellates (Protoperidinium sp.,Amphisoleta sp. and Gymnodinium sp.). Instantaneous growth ratesof nanophytoplankton (<20 µm) varied between 0.019and 0.080 day^–1, equivalent to between 0.03 and 0.12 chlorophylldoublings day^–1. Instantaneous grazing rates of microzooplanktonon nanophytoplankton varied from 0.012 to 0.052 day^–1.This corresponds to a nanophytoplankton daily loss of between1.3 and 7.0% (mean = 3.76%) of the initial standing stock, andbetween 45 and 97% (mean = 70.37%) of the daily potential production.Growth rates of microphytoplankton (>20 µm) were lower,varying between 0.011 and 0.070 day^–1, equivalent to 0.015–0.097chlorophyll doublings day^–1. At only three of the 10 stationsdid grazing by microzooplankton result in a decrease in microphytoplanktonconcentration. At these stations instantaneous grazing ratesof microzooplankton on microphytoplankton ranged between 0.009and 0.015 day^–1, equivalent to a daily loss of <1.56%(mean = 1.11%) of initial standing stock and <40% (mean =28.55%) of the potential production. Time series grazing experimentsconducted at 6 h intervals did not show any diel patterns ofgrazing by microzooplankton. Our data show that microzooplanktongrazing at the ice edge were not sufficient to prevent chlorophylla accumulation in regions dominated by rnicrophytoplankton.Here, the major biological routes for the uptake of carbon thereforeappear to be grazing by metazoans or the sedimentation of phytoplanktoncells. Under these conditions, the biological pump will be relativelyefficient in the drawdown of atmospheric CO₂.     

Sinking rates of heterogeneous, temperate phytoplankton populations

Throughout the summer of 1978, the sinking rates of phytoplanktonwithin the Controlled Experimental Ecosystems (CEE's) were monitoredusing a technique based upon measurement of the transit timeof radioactively (¹⁴C) labeled cells. The experimental frameworkof FOODWEB 1 offered an unprecedented opportunity to documentthe sinking rates of heterogeneous phytoplankton of diversetaxonomic composition, growing under a variety of nutrient regimes;the absence of advective exchange in the CEE's provided knowledgeof the preconditioning history of the phytoplankton sampledat any given time. Sinking rates of whole phytoplankton assemblages (not size-fractioned)ranged from 0.32 – 1.69 m·day^–1; the averagerate (± s.d.) observed was 0.64 ± 0.31 m·day^–1.The most notable deviations from the mean value occurred whenthe population size distribution and taxonomic composition shifteddue to blooms. The relationship between phytoplankton sinkingand ambient nutrient levels was studied by following the ratesof a given size fraction (8–53 µm) for ten daysfollowing nutrient enrichment of a CEE. Over this time sinkingrates ranged from 1.08– 1.53 m·day^–1; decreasedrates occurred after nutrification, yet over the course of theentire trial sinking rates were not significantly (p >0.05)correlated to the ambient levels of any single nutrient species. The sinking rate implications of spore formation were also studied,and showed that sinking rates of Chaetoceros constrictus andC. socialis spores (2.75 ± 0.61 m·day^–1)were ca 5-fold greater than rates measured when the vegetativestages of these species dominated the population, reflectingthe influence of physiological mechanisms in controlling phytoplanktonbuoyancy. An example of the potential influence of colony formation uponbuoyancy was noted in observations of C. socialis which occasionallywas found to exist in large spherical configurations made ofcoiled cell chains and having low (0.40 m·day^–1)sinking rates. A hydrodynamic rationale is presented to showhow such a colony together with enveloped water may behave asa unit particle having lower excess density, and therefore lowobserved sinking rate, despite its conspicuously large size. Overall, sinking rates were not significantly correlated withany of the measured nutrient or photic parameters. There were,however, trials and comparisons which showed evidence for: (1)higher sinking rates in populations dominated by large cells,(2) decreased sinking rates after nutrient enrichment, and (3)buoyancy response to light levels. It is suggested that correlationof sinking rates with synoptic environmental measurements atany given time is not explicit because the associations mayinvoke lag times of physiological response. The interpretationmade from these findings is that the preconditioning historyof the population, rather than the prevailing conditions atthe time of a given measurement, is most closely associatedwith population buoyancy modifications.     

Seasonal nitrogen assimilation and carbon fixation in a fjordic sea loch

Carbon (C) fixation and nitrogen (N) assimilation rates havebeen estimated from ¹⁴C and ¹⁵N techniques for a 12 month periodin a Scottish sea loch. The maximum rate of nitrogen assimilated(29.92 mmol N m^–2 day^–1) was in April at the mostseaward station; similar high rates were experienced duringMay at the other stations. Carbon fixation rates were maximal(488–4047 mg C m^–2day^–1) at the time of highphytoplankton biomass (maximum 8.3 mg m^–3 chlorophylla) during May, whilst nitrate concentrations remained >0.7µ.mol l^–1. C:N assimilation ratios suggest nitrogenlimitation only during the peak of the spring bloom, althoughat times nitrogen (nitrate and ammonium) concentration fellto 0.2 µmol l^–1 in the following months. The verticalstability of the water column, influenced by tidal and riverineflushing, varied along the axis of the loch, resulting in markeddifferences between sampling stations. Although ammonium waspreferentially assimilated by phytoplankton, >50% of productionwas supported by nitrate uptake and only during the summer monthswas the assimilation of ammonium quantitatively important.     

Biomass, feeding and production of Noctiluca scintillans in the Seto Inland Sea, Japan

The abundance and biomass of the large heterotrophic dinoflagellateNoctiluca scintillans, together with the changes in its potentialprey items, were monitored in the Seto Inland Sea, Japan, duringsummer 1997 (17 July-11 August). Growth and grazing rates ofNscintillans fed natural plankton populations were also measuredeight and seven times, respectively, during the survey period.The abundance and biomass of N scintillans averaged over thewater column (19 m) were in the range 1–345 cells 1^–1(temporalaverage = 93 cell1^–1) and 0.1–49.6 µg C l^–1(temporalaverage = 13.8 µg C l^–1; three times higher thanthat of calanoid copepods during the same period). Noctilucascintillans populations followed the changes in phytoplankton:N.scintillans biomass was increasing during the period of diatomblooms and was at a plateau or decreasing during periods oflow chlorophyll a. The growth rates of N.scintillans (µ)were also consistent with the wax and wane of the N.scintillanspopulation: N.scintillans showed highest growth rates duringdiatom blooms. A simple relationship between µ and chlorophylla concentration was established, and the production of N.scintillanswas estimated using this relationship and the measured biomass.The estimated production averaged over the water column wasin the range >0.1–5.2 µg C l^–1 day^–1(temporalaverage = 1.4 µg C l^–1 day^–1; 64% of the productionof calanoid copepods during the same period). Diatom clearancerates by N.scintillans were in the range 0.10–0.35 mlcell^–1 day^–1, and the phytoplankton population clearanceby N.scintillans was >12% day^–1. Thus, although thefeeding pressure of N.scintillans on phytoplankton standingstock was low, N.scintillans was an important member of themesozooplank-ton in terms of biomass and production in the SetoInland Sea during summer.     

Winter-spring variability of size-fractioned autotrophic biomass in Concepcion Bay, Chile

The temporal variability of size-fractioned autotrophic biomassat three depth levels (1, 8 and 25 m) was studied during thewinter-spring transition at two oceanographic stations in ConcepciónBay. Size spectra were obtained on eight occasions by two differentmethods: (i) determining the biomass of seven autotrophic sizefractions by in vivo fluorescence; and (ii) measuring the filamentlength of chain-forming diatoms through direct microscopy. Aclear vertical gradient of biomass was found in all profiles,with maximum values in the surface layer (1 and 8 m levels).Values of chlorophyll were on average 6.2 (range 1.08–25.67)times higher at 1 m than at 25 m, and 7.4 (range 1.15–26.83)times more at 8 m than at 25 m. On a temporal basis, total biomassincreased from low average values in winter (2.5 mg chl-a m^–3)to high values in late spring (11.6 mg chl-a m^–3). Duringthe whole sampling period (June 8-November 19), the nano- andnet-plankton (1.8–40 µm and 40–335 µmsize fractions respectively) were more abundant near the surface(1 and 8 m depth) than close to the bottom (25 m depth); however,the picoplankton fraction (<1.8 (µm) showed an inverserelationship, with a slight trend to increase near the bottomtoward spring. The highest absolute biomass was concentratedin the net-plankton fraction during the whole period and therelative importance of the picoplankton decreased from winter(6.50 and 15.5% for shallow and bottom levels) to spring (1.5and 10.3% for shallow and bottom levels). This relative effectis caused by the higher absolute values of biomass observedin the net-plankton fraction toward spring. These changing patternsshould have an impact in the size-composition and abundanceof higher trophic levels, mainly through grazing, in particularby modifying food availability to microfJagellates, ciliatesand filter-feeding zooplankton.     

Diel variability in nitrogenous nutrient uptake by phytoplankton in the Chesapeake Bay plume

Diel patterns in the uptake of nitrogenous nutrients were observedin the coastal plume of the Chesapeake Bay system, but the specificpatterns varied with season. During the winter months, ratesof NH₄⁺ and urea uptake were significantly higher at night thanduring the day, and rates of NO₃^– uptake were higher duringthe day. During the summer, rates of NH₄⁺ and urea uptake weresignificantly higher at night only during half the studies conducted;during the remaining studies, there was either no significantdifference or rates of uptake of NH₄⁺ were higher during theday. Rates of NO₃^– uptake during the summer months werealso higher during the day than at night. Seasonal differenceswere also apparent in the time of day at which maximum observeduptake rates of each nitrogen nutrient occurred. During thewinter-spring months, maximum observed rates of NO₃^– uptakeoccurred between first light and noon, whereas during the summermonths, maximum observed uptake rates of NO₃^– occurredboth morning and afternoon, and consistently 9–16 h afterthe maximum observed peak in the uptake of reduced nitrogen.We interpret these findings in terms of seasonal shifts in nitrogennutritional status of the assemblages, as well as species-specificdifferences in the effect of a given stimulus (e.g. a nitrogenpulse at the mouth of the Bay) to entrain an uptake response,and we suggest that the extent of this variability must be understoodbefore generalizations about the use of f-ratios as characteristicsof specific populations or water masses can be drawn.