Diurnal variations in the contributions of autotrophic picoalgae and heterotrophic bacteria to planktonic production in an upland lake

An investigation of the diurnal variation in contributions toproduction of the autotrophic and heterotrophic components ofthe picoplankton community was carried out during August andSeptember in Llyn Padarn, a mesotrophic upland lake in NorthWales. The picoplankton was separated using 1 µm pore-sizedfilters into the smaller cell sized fraction (<1 µm),the majority of the bacteria and the larger cell sized picoalgae(<3>1 µm), together with some bacteria. The distributionof bacterial heterotrophic activity between these two fractionsof picoplankton was assessed by uptake of [¹⁴C]glucose and differentialfiltration. Thus, the absolute autotrophic production by picoalgaeand the heterotrophic contribution by bacteria to picoplanktoncommunity production via uptake of extracellular organic carbon(EOC) were determined. Rates of picoplankton community productionexhibited diurnal variation with maximum rates of 19.1 mg Cm^–3 h^–1 recorded at 18.00 h at 4 m depth in September.The bacterial contribution to picoplankton community productionincreased markedly between 15.00 and 18.00 h. Rates of absoluteautotrophic production varied less over 24 h than rates of accumulationin bacteria of ¹⁴C-labelled EOC released from the entire phytoplanktoncommunity. Bacteria contributed up to 86–98% of the neworganic carbon within the picoplankton community at the endof the day. The maximum rate of absolute autotrophic productionin the picoplankton was 1.6 mg C m^–3 h^–1 at 18.00h at 1 m in August, and the maximum rate of bacterial accumulationof new organic carbon was 18.5 mg C m^–3 h^–1 at 18.00h in September at 4 m depth. The diurnal pattern of picoplanktoncommunity production involved increasing rates during the daywith a maximum at 18.00 h. Autotrophic processes were dominantin the first 3–6 h of the light cycle and heterotrophicuptake of ¹⁴C-labelled EOC was the major component from 15.00h onwards. Bacterial uptake of newly released EOC by phytoplanktonwas rapid, comprised the majority of picoplankton production,particularly later in the day, and contributed a maximum of60% of the total pariculate primary production in plankton between15.00 and 18.00 h at 4 m in September with a mean contributionof between 6 and 26% over 24 h in these investigations. Theimportance of autotrophic processes in picoplankton communityproduction has been overestimated in previous investigations.Bacteria play a major role in transferring newly produced EOCrapidly from phytoplankton to the picoplankton community. Atthe end of the day, the majority of newly produced organic carbonis in bacterial cells and this production is significant inthe dynamics of carbon production within the entire planktoniccommunity.     

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 (     

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.     

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.     

Autotrophic picoplankton in southern Lake Baikal: abundance, growth and grazing mortality during summer

Autotrophic picoplankton were highly abundant during the thermalstratification period in late July in the pelagic area (waterdepth 500–1300 m) of southern Lake Baikal; maximum numberswere 2 x 10⁶ cells ml^–1 in the euphotic zone ({small tilde}15m). Unicellular cyanobacteria generally dominated the picoplanktoncommunity, although unidentified picoplankton that fluorescedred under blue excitation were also abundant (maximum numbers4 x 10⁵ cells ml^–1) and contributed up to {small tilde}40%of the total autotrophic picoplankton on occasions. Carbon andnitrogen biomasses of autotrophic picoplankton estimated byconversion from biovolumes were 14–84 µg C l^–1and 3.6–21 µg N l^–1. These were comparableto or exceeded the biomass of heterotrophic bacteria. Autotropicpicoplankton and bacteria accounted for as much as 33% of paniculateorganic carbon and 81% of nitrogen in the euphotic zone. Measurementsof the photosynthetic uptake of [^l4C]bicarbonate and the growthof picoplankton in diluted or size-fractionated waters revealedthat 80% of total primary production was due to picoplankton,and that much of this production was consumed by grazers inthe <20 µ.m cell-size category. These results suggestthat picoplankton-protozoan trophic coupling is important inthe pelagic food web and biogeochemical cycling of Lake Baikalduring summer.     

Summer nutrient dynamics of the Middle Atlantic Bight: nitrogen uptake and regeneration

Nitrate and ammonium uptake and ammonium regeneration rates(by zooplankton, microplankton and benthos) were measured onthe Atlantic continental shelf (Middle Atlantic Bight) duringsummer, 1980. Euphotic zone profiles of NO₃^– and NH₄⁺uptake rates were similar in magnitude and vertical structureover a large geographical area. Microplankton NH₄⁺ regenerationrates, although measured less frequently, also showed a relativelyconsistent vertical structure; rates were positively correlatedwith uptake rates. Nitrate assimilation (‘new’ production)was used to estimate vertical eddy diffusivity and paniculatesinking rates. Eddy diffusion estimates ranged from <0.1to >2.0 cm² s^–1 and were positively related to arealprimary production. Estimated particulate sinking rates averaged5 mg at Nm^–2d^–1 and compared favorably with sedimentationrates measured from free-floating and moored sediment traps.Benthic nitrogen regeneration rates represented <10% of thispaniculate nitrogen flux. Within the mixed layer, NH₄⁺ assimilation(‘regenerated’ production) represented 50–80%of the total (NO₃^– + NH₄⁺ ) nitrogen productivity and33% for the euphotic zone. Of this, 30% was attributed to zooplankton,63% to microplankton (<100 µm) and 7% to benthos. Onthe average, 74% of the microplankton NH₄⁺ regeneration wasassociated with organisms passing 1 µm filters.     

Partitioning of the food ration of marine ciliates between pico- and nanoplankton

Food size-range for 13 species of Tintinnina and 18 species of Oligotrichina were studied using electronic particle counting and in situ observation of food vacuole contents. Tintinnids consume nanoplankton in the size range 2–20 μm. Oligotrichous naked ciliates consume particles in the size range 0.5–10 μm. Ciliates smaller than 30 μm take 72% picoplankton and 28% nanoplankton. For ciliates between 30 μm and 50 μm the proportions are reversed (30% pico- and 70% nanoplankton), while the larger ciliates (> 50 μm) take nanoplankton almost exclusively (95% nano- and 5% picoplankton). A seasonal study of total Oligotrichida grazing showed that natural particles were consumed at rates that varied from 1 to 20 μg C 1⁻¹ day⁻¹. This included between 1 and 38% of the bacterioplankton production and 9 to 52% of the nanoplankton production. In the N-W Mediterranean the total ciliate production varied from 0.4 to 8.2 μg C 1⁻¹ day⁻¹. Research supported by CNRS-PIROCEAN AIP-RTM-953146-GRECO P4 and by CNRS-UA 716 (FR), and by the University of Nice and by the CNRS-GRECO 88 (MLP)     

Microplankton and its function in a zone of shallow hydrothermal activity:the Craternaya Bay, Kurile Islands

Biomass and productivity of microplankton were measured in theCraternaya Bay (Kurile Islands), which is influenced by hydrothermalactivity and volcanic heating. The hydrothermal fields are situatedaround its shores and underwater within the 0–20 m depth.A dense ‘bloom’ of photoautotrophic microplanktonwas observed there, dominated by diatoms, phytoflagellates andthe symbiont-containing ciliate Mesodinium rubrum. The biomassof these ciliates attained 3–11 g m^-3 in the upper waterlayer. The total biomass of the phototrophic microplankton reached30–46 g m^-3. The primary production in the water columnwas, correspondingly, enormously high: 6–10 g C m^-2 day^-1.The depth of the euphotic zone was 7 m. Pelagic photosynthesiswas inhibited in the upper 0–1 m by the spreading of alayer of low-salinity hydrothermal water. The numerical densityof bacterioplankton in the upper zone of the water column variedfrom 1 x 10⁶ to 2.9 x 10⁶ cells ml^-1, and its wet biomass from250 to 750 mg m^-3. Its production varied at stations from 70to 390 mg m^-3 day^-1. Chemosynthesis contributed up to 30% ofthis production in the sites neighbouring the hydrothermal vents.Outside their direct impact however, its share was negligible.The biomass of heterotrophic planktonic ciliates varied from30 to 270 mg m^-3. The mechanisms of possible influence of shallowvolcanic activity on development and function of microplanktonin the Craternaya Bay is discussed.     

Bacterial growth and grazing loss in contrasting areas of North and South Atlantic

Samples were collected from the top 200 m of the water columnat 50 stations during two cruises in different, near equinoctialseasons on an Atlantic transect near the 20°W meridian between50°N and 50°S. These samples were analysed to determinecharacteristics of the heterotrophic bacterial populations.Flow cytometry was used to enumerate these bacteria and determinetheir average size so as to calculate their biomass. Heterotrophicbacterial production, and the rate of grazing of these bacteriaby heterotrophic nanoplankton in the main depth layers, weredetermined using ³H thymidine and ¹⁴C leucine techniques. Thebiomass of heterotrophic nanoplankton in these layers was determinedusing a glucosaminidase assay. Five provinces were distinguishedalong the transect and characterized by average values of allmeasured parameters. The relative composition and activity ofthe microbial community in the water columns within each provincechanged little between the two cruises. Lowest heterotrophicbacterial biomass of 1–2 mg C m^–3 and productionof 0.1–0.2 mg C m^–3 day^–1 were found in thenorthern and southern Atlantic gyres, and were relatively similarin both seasons. Biomass and production were 2–4 timeshigher in the northern and southern temperate waters, and inequatorial waters, than in the gyres and tended to show moreseasonal variation. Production and biomass in the layer belowthe pycnocline were lower by 10–30% and about 50%, respectively,than values determined in the surface mixed layer, and variedless with latitude. Depth-integrated values of these two parameterswere generally of similar size in the mixed water layer andthe layer of the chlorophyll maximum and pycnocline, and tendedto vary with season. The specific growth rate of heterotrophicbacteria was in the range 0.05 to 0.12 day^–1 in the topmixed layer at all latitudes. In spite of the elevated temperatures,bacterial growth appears to be restricted by a shortage of nutrientsso that the microbial community cycles very slowly, with a turnovertime of the order of 1 week or more. The depth-integrated biomassof heterotrophic nanoplankton was generally about 100% of theheterotrophic bacterial biomass in the same water. Grazing bythese nanoplankton at the rate measured could consume all ofthe new production of heterotrophic bacteria in all waters,and they probably control the populations of both heterotrophicand phototrophic bacteria.     

Contribution of Picocyanobacteria to total primary production and community respiratory losses in a backwater system

The contribution of autotrophic picoplankton (APP) to phytoplanktonicprimary production, investigated during the phytoplankton growingseason (March–September) in a macrophyte-dominated backwatersystem near Vienna, showed that APP mainly consisted of rod-shapedand coccoid cyanobacteria. Two stations were examined, exhibitingsimilar seasonal patterns in the development of picocyanobacteria,although the two sites differed in picocyanobacterial cell numbersand biomass by a factor of 1.5. Cell numbers determined by epifluorescencemicroscopy varied between 0.29 x 10⁴ and 34.5 x 10⁴ cells ml^–1at Station 1, and between 0.23 x 10⁴ and 19.1 x 10⁴ cells ml^–1at Station 2. At both sites, the mean cell volume of picocyanobacteriawas 0.5 µm³. Carbon fixation in the planktonic communityof the Kühwörter Wasser was dominated primarily bylarger phytoplankton, although the picoplankton community sometimessupplied up to 74% (mean: 35%) of total primary production.Distinct differences in chlorophyll a concentrations and primaryproduction between the two sites refer to a greater competitionbetween phytoplankton and macrophytes at Station 2. Communityrespiration deviated greatly in time and in level at the twostations, showing a higher dynamic in community metabolism atStation 1. At this site, community respiration losses rangedbetween 12 and 100% of gross production. Hence, community metabolismcomprised net autotrophic, balanced, and net heterotrophic situationsover the investigation period, whereas at Station 2, only netautotrophic situations could be determined.     

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.     

The importance of phytoflagellate, heterotrophic flagellate and ciliate grazing on bacteria and picophytoplankton sized prey in a coastal marine environment

The uptake of bacteria and picoplankton sized fluorescentlylabelled beads was measured off the west coast of the SouthIsland of New Zealand in winter. Phytoflagellates and heterotrophicflagellates showed similar grazing rates on 0.49 µm beads,with mean clearance rates of 1.1 and 1.8 nl ind.^–1 h^–1,respectively. Clearance rates for 1.09 µm beads were 0.9nl ind.^–1 h^–1 for heterotrophic flagellates and0.5 nl ind.^–1 h^–1 for phytoflagellates. Non-loricateciliates had clearance rates of 1.5 µl ind.^–1 h^–1for the picoplankton sized particles. The heterotrophic flagellatesshowed no significant difference between clearance rates of0.49 and 1.09 µm particles. Phytoflagellates, however,showed an apparent preference for the smaller particles. Themeasurement of significant grazing by phytoflagellate populationsin the marine environment is important and indicates that weneed to reassess our concepts of food web structure.     

Production of new organic carbon and its distribution between autotrophic picoplankton, bacteria, extracellular organic carbon and phytoplankton in an upland lake

• 1 Picoplankton community production (0.2–2μm) was investigated over 3 months, June-September 1991, in Llyn Padarn, a mesotrophic upland lake in north Wales.
• 2 The picoplankton was differentiated into autotrophic algae (<1–3μm) and heterotrophic bacteria (<0.2–1 μm) using differential filtration through a 1 μm pore size Nuclepore filter.
• 3 Efficient separation of these distinct metabolic constituents of picoplankton was obtained. A good correlation (r= 0.81, P < 0.001) was found between physical separation of bacterial and picoalgal cells from fluorescence microscopy and the distribution of heterotrophic metabolic activity between different cell size fractions measured by uptake of ¹⁴C-glucose.
• 4 Picoplankton community production was differentiated into the ‘absolute’ autotrophic production by picoalgae, corrected for overestimation due to retention of bacteria with the picoalgae, and the heterotrophic component, bacterial uptake of ‘extracellular organic carbon’ (EOC), derived from the entire phytoplankton community.
• 5 The heterotrophic contribution to picoplankton community production ranged from 88 to 1%, mean value 55% of total. Autotrophic picoplankton production was dominant in June and July, but in August and September heterotrophic uptake of EOC was the major input to picoplankton community production.
• 6 During the 3 months, the mean contributions to plankton production were autotrophic picoplankton 10.3%, heterotrophic bacterial uptake of EOC 9.7%, EOC in lake water 11.6% and phytoplankton (>3μm) 68.3%.
• 7 Bacteria accounted for about half the picopfankton community production via uptake of EOC. Thus although autotrophic picoplankton were ubiquitous, it is likely that their contribution via primary production to the carbon balance of planktonic environments has been overestimated in previous studies.

     

Mesozooplankton grazing under conditions of extreme eutrophication in Guanabara Bay, Brazil

The objective of the present study was to quantify mesozooplanktongrazing in the eutrophic waters of Guanabara Bay. Mesozooplankton(>200 µm) was dominated by the copepods Acartia lilljeborgi,Acartia tonsa, Parvocalanus crassirostris and Paracalanus furcatus.Dinoflagellates, specifically the species Prorocentrum triestinum,were an important group for mesozooplankton nutrition, beingingested in significant amounts during all experiments. On average,12.3 ± 2.9 P. triestinum cells were ingested copepod^–1min^–1 (other dinoflagellates: 11 ± 8 cells copepod^–1min^–1). Filamentous cyanophyceae and nanoplankton wereingested in one experiment each, but the mesozooplankton communitygenerally preferred dinoflagellates to these groups, which werealways abundant in the water column. Euglenophyceae were notingested, although they dominated in one experiment. Mesozooplanktoningested, on average, only 0.2% of the nano- and microplanktonbiomass per day. The results suggest that grazing was not acontrolling process for the nano- and microplankton communityin the study area. Addition of zoeae larvae of Chasmagnatusgranulata (Decapoda: Brachyura: Grapsidae) in one experimenthad a significant effect on the mortality of adult copepods,probably due to a predator–prey relationship.     

Nitrogen dynamics in lower Narragansett Bay, Rhode Island. I. Uptake by size-fractionated phytoplankton populations

The contribution of nanoplankton (< 10 µm fraction)to winter – spring (1977 – 78) and summer (1978,1979) phytoplankton nitrogen dynamics in lower NarragansettBay was estimated from ammonium, nitrate and urea uptake ratesmeasured by ¹⁵N tracer methods. During the winter – spring,an average of 80% of chlorophyll a and nitrogen uptake was associatedwith phytoplankton retained by a 10 µm screen. In contrast,means of 51 – 58% of the summer chlorophyll a standingcrops and 64 – 70% of nitrogen uptake were associatedwith cells passing a 10 µm screen. Specific uptake ratesof winter – spring nanoplankton populations were consistentlylower than those of the total population. Specific uptake ratesof fractionated and unfractionated summer populations were notsignificantly different. Ammonium uptake averaged between 50and 67% of the total nitrogen uptake for both the total populationand the < 10µm fraction. The total population and the10 µm fraction displayed similar preferences for individualnitrogen species. Though composed of smaller cells, flagellatedominated nanoplankton assemblages may not necessarily takeup nitrogen at faster rates than diatom dominated assemblagesof larger phytoplankters in natural populations. ¹Present address: Australian Institute of Marine Science, P.M.B.No. 3, Townsville M.S.O., Qld. 4810, Australia     

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.     

Abundance and productivity of heterotrophic nanoplankton in Georgia coastal waters

The population abundances and rates of biomass production ofheterotrophic nanoplankton (HNAN) in Georgia coastal waterswere evaluated by epifluorescence microscopy. HNAN populations(mostly non-pigmented microflagellates <10 µm in diameter)ranged from 0.3 x 10³ cells ml^–1 in shelf waters 15 kmoffshore to 6.3 x 10³ cells ml^–1 in waters 0.25 km fromthe coast. There was a strong correlation (r = 0.83) betweenHNAN and free bacterioplankton population abundances, but noapparent relation (r = 0.38) between HNAN and phototrophic nanopLankton(PNAN) abundances. HNAN biomass production in estuarine andnearshore shelf waters, as estimated from increases in HNANpopulations during laboratory incubations of natural water samples,ranged from 0.10 to 0.79 mg C m^–3 h^–3, with populationgeneration times of 9.7 to 26.5 h. There was a significant linearrelation (r = 0.95) between HNAN biomass and HNAN productivity.We calculated that HNAN may graze at least 30% to 50% of dailybacterioplankton production in Georgia coastal waters.     

Feeding and reproduction of Calanus finmarchicus during non-bloom conditions in the Irminger Sea

Simultaneous ingestion and egg production experiments were conductedwith female Calanus finmarchicus in April/May and July/August2002 in the Irminger Sea. Experimental animals were providedwith natural microplankton food assemblages and incubated underin situ conditions for 24 h. The quantity of food consumed wassignificantly related to the concentration of prey cells, withtotal daily ingestion rates ranging from 0.6 to 8.1 µgof carbon female^–1 day^–1, corresponding to carbon-specificrates of 0.6–4.7% day^–1. Egg production rates (EPRs)remained relatively low (0.3–11 eggs female^–1 day^–1)during both periods of investigation and were not influencedby food availability. The data were used to construct energeticbudgets in which the microplankton carbon ingested, includingciliates, was compared with the carbon utilized for egg productionand respiration. These budgets showed that ingestion alone couldnot provide the necessary carbon to sustain the observed demandsfor growth and metabolism. Although ciliates constituted >80%of the total material ingested at times, they were not sufficientto provide the metabolic shortfall. Indeed, the females weretypically lacking 5 µg of carbon each day, 5% of theircarbon biomass. Our study results highlight the possible importanceof internal reserves in sustaining reproduction in C. finmarchicusduring periods of food scarcity.     

Spatial Distribution, Structure, Biomass, and Physiology of Microbial Assemblages across the Southern Ocean Frontal Zones during the Late Austral Winter

We examined the spatial distributions of picoplankton, nanoplankton, and microplankton biomass and physiological state relative to the hydrography of the Southern Ocean along 90 degrees W longitude and across the Drake Passage in the late austral winter. The eastern South Pacific Ocean showed some large-scale biogeographical differences and size class variability. Microbial ATP biomass was greatest in euphotic surface waters. The horizontal distributions of microbial biomass and physiological state (adenylate energy charge ratio) coincided with internal currents (fronts) of the Antarctic Circumpolar Current. In the Drake Passage, the biological scales in the euphotic and aphotic zones were complex, and ATP, total adenylate, and adenylate energy charge ratio isopleths were compressed due to the extension of the sea ice from Antarctica and constriction of the Circumpolar Current through the narrow passage. The physiological state of microbial assemblages and biomass were much higher in the Drake Passage than in the eastern South Pacific Ocean. The temperature of Antarctic waters, not dissolved organic carbon, was the major variable controlling picoplankton growth. Estimates of picoplankton production based on ATP increments with time suggest that production under reduced predation pressure was 1 to 10 mug of carbon per liter per day. Our results demonstrate the influence of large-scale hydrographic processes on the distribution and structure of microplankton, nanoplankton, and picoplankton across the Southern Ocean.     

Zooplankton growth rates: the larvaceans Appendicularia, Fritillaria and Oikopleura in tropical waters

The growth rates of Appendicularia sicula, Fritillaria borealissargassi, Fritillaria haplostoma, Oikopleura dioica and Oikopleuralongicauda were determined from microcosms incubated in situat 23C in Jamaican waters. Experiments were conducted fromoligotrophic offshore waters, through mesotrophic Lime Cay andeutrophic Kingston Harbour in both natural and nutrient-enhancedphytoplankton communities. Length-weight relationships werecalculated for two of these species: O.longicauda log W=2.47log TL –6.10 and F.haplostoma log W=2.44 log TL –7.37,where weight (W) is in micrograms and trunk length (TL) is inmicrometres. Instantaneous growth rates averaged 1.7–2.5day^–1 for the five species and were observed as high as3.3 day^–1 These instantaneous rates are equivalent todaily specific growth rates averaging 4.6–11.4 and rangingup to 28. In larger genera, growth rates were related positivelyto picoplankton and nanoplankton concentration, and negativelyto the biomass of larvaceans, but in the smallest species growthwas unrelated to these factors. However, because the variabilityin these two factors within microcosms exceeded their naturalrange of variability, growth rates of larvaceans may normallybe unlimited by resources or population density effects. ¹Present address :Monterey Bay Aquarium Research Institute 7700Sandholdt Road, Moss Landing, CA 95039-0628, USA ²Present address :Bedford Institute of Oceanography PO Box 1006,Dartmouth, Nova Scotia B2Y 4A2, Canada