Nitrogen uptake by heterotrophic bacteria and phytoplankton in Arctic surface waters

We estimated rates of heterotrophic bacterial and phytoplanktonuptake of nitrate, ammonium, and urea using ¹⁵N-labelled nitrogenand specific metabolic inhibitors of prokaryote and eukaryotenitrogen metabolism in the surface waters of the North Water(northern Baffin Bay) during autumn that were characterizedby the absence of cyanobacteria (comprising prochlorophytes).The percentage of nitrate + ammonium uptake by heterotrophicbacteria ranged between 44 and 78% of the measured total uptakeand was the highest when the phytoplankton biomass was relativelylow (<2 µg Chlorophyll a L^–1). Phytoplanktonaccounted for a larger fraction (e.g., 58–95%) of ureauptake than heterotrophic bacteria. When our results are combinedwith those from previous studies carried out in diverse temperateand polar areas, it appears that heterotrophic bacteria accountfor 25% (14–40%; median and interquartile range) of thetotal nitrate uptake in surface waters with chlorophyll biomass<2 µg L^–1. Estimates of new production computedfrom phytoplankton carbon uptake and f-ratios may be stronglyoverestimated in regions where nitrate uptake by heterotrophicbacteria is high and the biomass of phytoplankton is low.     

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.     

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     

Bacterial response to seasonal changes in primary production and phytoplankton biomass in Lake Constance

Parameters characterizing bacterial biomass and metabolic activityare compared with phytoplankton biomass and daily primary productionrates throughout the year. Between late March (before the onsetof the phytoplankton spring bloom) and mid-July (diatom maximum),bacterial degradation of organic matter was more closely relatedto phytoplankton productivity than during the rest of the year.Bacterial production (as estimated by amino acid net uptake)was significantly correlated with concentrations of chlorophyll and pheopigments. However, bacterial production was correlatedless closely with primary production and only weakly with bacterialbiomass. Bacterial biomass was also only weakly correlated withprimary production but significantly with pheopigments. Numbersof active bacteria as estimated by autoradiography covariedclosely with bacterial production and cell numbers. Wheneverbacterial production was low, enhanced proportions of aminoacids were respired. Oxygen consumption measurements showedthat the size fraction <3 µm contributed 25–75%to total respiration. On average, bacterial biomass comprised11 % of paniculate organic matter and roughly equalled phytoplanktonbiomass. During the growing season, bacterial production inthe uppermost 20 m comprised about 20% of phytoplankton productionwith large seasonal fluctuations. A tentative carbon budgetof the euphotic zone including primary production, zooplanktongrazing, bacterial production and sedimentation is presented. ¹Present address: Institute of Marine Resources A-018, ScrippsInstitution of Oceanography, University of California, San Diego,La Jolla, CA 92093, USA     

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 uptake and regeneration of inorganic nitrogen and phosphorus in a large oligotrophic lake: size-fractionation and antibiotic treatment

Uptake and regeneration of inorganic N and P in oligotrophicFlathead Lake (Montana) were measured with ¹⁵N and ³²P incorporationand dilution experiments, six times over a seasonal cycle. Theannual mean molar N P uptake ratio at ambient concentrationswas 13 9 (range = 4 8–34.2); uptake of nitrate, ammoniumand phosphate were always below saturation indicating both Nand P deficiency Organisms >280 µm were responsiblefor 0–60% of ammonium and 0–40% of phosphate regeneration,40–100% of the ammonium and 34–98% of phosphateregeneration occurred in the <3 µm fraction The <3µm fraction accounted for 7–70% of the ammoniumand 6–64% of the phosphate uptake. Results from antibiotictreatments indicated that both prokaryotic and eukaryotic ammoniumuptake was important, and that eukaryotes accounted for 53–98%of the ammonium regeneration. During thermal stratification,heterotrophic ammonium and phosphate regeneration by organisms>3 µm supplied much of the inorganic N and P in theepilimnion. Estimated rates of allochthonous and diffusive (i.e‘new’) ammonium, nitrate and phosphate input were<5% of biotic regeneration. These results suggests that (i)both N and P dynamics should be considered when examining nutrientregulation of primary productivity of oligotrophic lakes, (ii)bacteria probably compete with phytoplankton for both ammoniumand phosphate, (iii) biotic regeneration is the main sourceof nutrients to the epilimnion during stratification, and (iv)crustacean zooplankton were relatively unimportant sources ofregenerated ammonium and phosphate.     

Grazing of copepod assemblages in the north-east Atlantic: the importance of the small size fraction

The North Atlantic was the site for the 1989 JGOFS Pilot Study,an international study of ocean fluxes in relation to the carboncycle. In this paper we present preliminary estimates of thegrazing pressure by copepod assemblages at four stations 60,56, 52 and 47°N, along the JGOFS 20°W transect, duringJune–July. Three major size fractions of mesoplanktoniccopepods were considered, small (200–500 µm), medium(500–1000 µm) and large (1000–2000 µm).At each station, copepod composition and abundance were analysedand the gut fluorescence method was used to estimate ingestionrates. The results support the importance of the small sizefraction relative to the other fractions, in terms of numericalabundance and their grazing impact. However, the total grazingpressure of copepods on phytoplankton was relatively minor duringthe period of sampling since the fraction of phytoplankton standingstock and primary production consumed by the copepods was onaverage <1 and 2% respectively. The implications of theseresults as well as the potential sources of bias involved inthese types of measurements and estimations are discussed.     

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.     

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.     

Food concentration and temperature effects on the demography of <Emphasis Type="Italic">Latonopsis</Emphasis> cf. <Emphasis Type="Italic">australis</Emphasis> Sars (Cladocera: Sididae)

This study has two main objectives, the first being the determination of net phytoplankton primary production to explain the phytoplankton’s function in a wetland carbon cycle, while the second objective is to relate this function with the phytoplankton assemblage composition. The annual variation in the phytoplankton production was monitored monthly for more than a year (2007–2008) in the semiarid eutrophic, hydrologically-perturbed “Tablas de Daimiel” National Park wetland. The phytoplankton fraction considered in this study comprised all organisms between the size 3 and 100 μm. The total biomass of phytoplankton was obtained by counting algae and calculating their volume, while net primary production and respiration were quantified by in situ incubations with the Winkler method. The respiration ranged from undetectable to 0.07 mgO₂ l⁻¹ h⁻¹; net photosynthesis reached 0.20 mgO₂ l⁻¹ h⁻¹. Net primary production was maximum at the end of the warm period (October 2007), and other peaks occurred at the start of summer (July 2007) or spring (March 2008). When maximum production took place, phytoplankton was mainly composed of small fast-growing chlorophytes (Tetraselmis cf. fontiana or Chlamydomonas sp.), in addition to some of the large, S-strategist algae (Peridinium umbonatum, Microcystis flos-aquae, Euglena sp.). The phytoplankton metabolism in “Tablas de Daimiel” was autotrophic as a whole due to changing contributions of algal groups. Only chlorophyte biomass was statistically related to net primary production. The conclusion reached is that this shallow eutrophic semiarid wetland possessed an annual net autotrophic production of phytoplankton fraction resulting from the small, fast-growing algae enhanced by hydrological perturbations that interrupted the autogenic course of S strategists.     

Uptake and regeneration of inorganic nitrogen in coastal waters influenced by the Mississippi River spatial and seasonal variations

The Mississippi and Atchafalaya Rivers introduce large amountsof nutrients to surface waters of the northern Gulf of Mexico.This paper reports the most complete data to date on inorganicnitrogen uptake and regeneration in a broad range of coastalenvironments influenced by the river water, along with informationon nutrient concentrations and including pico-, nano-, and microplanktonspecies composition. Nitrate in surface waters is greatly reducednear the river plume, at salinities between 5 and 25 PSU, wherethe largest variance in uptake rates was observed, and was coincidentwith peaks in surface chlorophyll. Despite the depletion ofnitrate, nitrogen limitation was a rare event during the study,because of relatively high ammonium concentrations (>1 µmolNH₄⁺ I^–1 and regeneration rates. Two contrasting situationscharacterize the seasonal nitrogen dynamics in surface shelfwaters. High nitrate input during the spring caused a largebloom in which the cells were well adapted to use nitrate.Thedominant phytoplankton species were chain forming diatoms, alsoreported in sediment-trap studies in the area. Ammonium regenerationonly accounted for a small fraction of the nitrogen requirementsduring the bloom. In contrast, the low flow of river water duringsummer resulted in low nitrate concentrations in surface water.In this case phytoplankton productivity was highly reduced andmay depend greatly on ‘in sita’ ammonium regeneration.     

Environmental control of phytoplankton assemblages in nearshore marine waters, with special emphasis on phototrophic ultraplankton

The present study assesses conditions that determine the dominanceof (i) phytoplankton by cells >5 µm versus cells <5p.m, (ii) phototrophic ultraplankton by prokaryotes versus eukaryotes,(iii) prokaryotes by phycocyanin-rich versus phycoerythrin-richcyanobacteria, and (iv) eukaryotes by cryptophyte versus non-cryptophytecells. In the Baie des Chaleurs, Gulf of St Lawrence, Canada,the decrease in phytoplankton <5 µm during the summeris related to nitrate and silicate exhaustion. In contrast,cells <5 µm are less affected by oligotrophic conditionsand are favoured by temperatures > 10°C. Ultraplanktoneukaryotes, because of larger cell volumes, may be selectivelygrazed by micro-zooplankton, and are likely to be more affectedby oligotrophic conditions than the dominating cyanobacteria.In the prokaryote assemblage, a relatively high contributionof phycocyanin-rich cyanobacteria is observed at times of lowsalinities and high attenuation of irradiance. Conversely, thedominant phycoerythrin-rich forms thrive in wanner and clearerwaters. Finally, the eukaryote assemblage is dominated by non-cryptophytecells, possibly because cryptophytes, which are larger in size,may be more affected by grazing than non-cryptophytes. The contributionof cryptophytes to the eukaryote assemblage increases at hightemperatures, high salinities and low nutrient concentrations.     

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.     

Size-fractionated productivity and nutrient dynamics of phytoplankton in subtropical coastal environments

It is now well established that the size distribution of phytoplankton plays an important role in primary production processes and nutrient dynamics of coastal environment. In situ observations showed that nanophytoplankton (3–20 μm) contributed 72.08% and58.18% of phytoplankton biomass and 58.32% and 41.14% of primary productivity to Xiamen Western Waters and the northern Taiwan Strait, respectively; picophytoplankton (0.2–3 μm) dominated the biomass (64.70%) and productivity (66.09%) in the southern Taiwan Strait. Furthermore, nanophytoplankton accounted for 75% of phosphate uptake with the highest rate constant (8.3×10^-5 s^-1) and uptake rate in unit water volume (5.4×10^-5 mmol dm^-3s^-1); picophytoplankton had the highest uptake rate in unit biomass (5.4×10^-5 mmol mg^-1s^-1) and photosynthetic index (3.8 mgC mgChl a^-1h^-1). All the results highlighted the remarkable characteristics of small size ranged (0.2–20 μm) phytoplankton in subtropical coastal environments: main contributor to phytoplankton biomass and production, high efficiency on organic carbon production and nutrient recycling. The far reaching environmental and ecological implications were discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Phytoplankton and zooplankton seasonal dynamics in a subtropical estuary: importance of cyanobacteria

A seasonal study of phytoplankton and zooplankton was conductedfrom 1999 to 2001 in Pensacola Bay, Florida, USA, to furtherthe understanding of pelagic food webs in sub-tropical estuaries.Monthly measurements included size-fractionated chlorophyll(whole water, <5 µm, <20 µm), net- and picophytoplanktoncomposition analyzed using microscopy, flow cytometry, and HPLCpigment analysis. Additionally, zooplankton abundance and dryweight were determined from net tows. The results show a phytoplanktoncommunity dominated by the small size fraction (<5 µm),especially during the warm periods. The <5 µm chlorophyllfraction was strongly correlated with cyanobacterial abundanceand zeaxanthin. Cyanobacteria (cf. Synechococcus) abundancepeaked during summer in the upper estuary, typically exceeding3 x 10⁹ L^-1, and was strongly correlated with temperature. Cyanobacteriaabundance at the freshwater end of the Bay (in the EscambiaRiver) was very low, suggesting that cyanobacteria were notdelivered via freshwater. Two pigmentation types of cyanobacteriawere observed. Phycoerythrin-containing cells (PE-rich) weremore abundant at the marine end, while phycocyanin-containingcells (PC-rich) were more abundant in the upper estuary. Thelarger algae (>5–10 µm) were predominantly composedof diatoms, followed by chlorophytes, cryptophytes and dinoflagellates.The three most abundant genera of diatoms were Thalassiosira,Pennales and Cyclotella. Zooplankton biomass averaged 12.2 µgC L^-1, with peak biomass occurring during May (     

Seasonal variation of phytoplankton community structure and nitrogen uptake regime in the Indian Sector of the Southern Ocean

This study investigates the dynamics of phytoplankton communities and nitrogen uptake in the Indian sector of the Southern Ocean during spring and summer. The study area is oligotrophic (Chl a stocks <50 mg m⁻²); nevertheless, a large spatial variation of phytoplankton biomass and community structure was observed. During both seasons the phytoplankton community in the seasonal ice zone showed higher biomasses and was mainly composed of large diatom cells. However, in the permanently open ocean zone the community had low biomass and was chiefly composed of nano- and picoflagellates. In the polar front zone, although biomass was higher, the community structure was similar to the open ocean zone. The results suggest that the variation in phytoplankton community structure on a larger scale resonates with gradients in water column stability and nutrient distribution. However, significant changes in biomass and nutrient stocks but little change in community structure were observed. Absolute nitrogen uptake rates were generally low, but their seasonal variations were highly significant. During spring the communities displayed high specific nitrate uptake (mean rate = 0.0048 h⁻¹), and diatoms (in the seasonal ice zone) as well as nano- and picoflagellates (in the permanently open ocean zone and polar front zone) were mainly based on new production (mean ƒ-ratio = 0.69). The transition to summer was accompanied by a significant reduction in nitrate uptake rate (0.0048 h⁻¹ → 0.0011 h⁻¹) and a shift from predominantly new to regenerated production (ƒ-ratio 0.69 → 0.39). Ammonium played a major role in the seasonal dynamics of phytoplankton nutrition. The results emphasize that, despite a large contrast in community structure, the seasonal dynamics of the nitrogen uptake regime and phytoplankton community structure in all three subsystems were similar. Additionally, this study supports our previous conclusion that the seasonal shift in nitrogen uptake regime can occur with, as well as without, marked changes in community structure. Received: 2 December 1997 / Accepted: 20 April 1998     

Respiration in eutrophic lakes: the contribution of bacterioplankton and bacterial growth yield

The contribution of bacterioplankton to total plankton respirationwas measured in two eutrophic Danish lakes and in experimentalenclosures treated with planktivorous fish and nutrients. Bacterialrespiration was calculated from measured oxygen uptake ratesin particles passing a 1.0-µm pore size filter, the rateswere then corrected for the size distribution of glucose uptake.During summer the respiration of the planktonic bacteria contributed{small tilde}50% of the community respiration in the two lakes.Prolific phytoplankton growth induced by biomanipulation andnutrient addition created situations where the contributionof the bacteria decreased to 20%. High bacterial contributionsto community respiration were found when the phytoplankton biomassdecreased. Simultaneous measurements of bacterial respirationand production (by means of [³H]thymidine incorporation) allowedan estimation of bacterial growth yield, which ranged from 9to 66%. In the two lakes the growth yield was constant witha mean of 29 ± 5% (±SD, RQ = 1). The variabilityof the growth yield was larger in the enclosures. The wide range(9–66%) was mainly caused by changes in bacterial netproduction without concomitant changes in respiration. The discussionincludes an evaluation of the oxygen uptake method in size fractionatedsamples and the availability of labile organic substrates asa factor controlling bacterial growth yield. Present address: Institute of Biology and Chemistry, Universityof Roskilde, P. Box 260, DK-4000 Roskilde, Denmark     

Nitrogen uptake by size-fractionated plankton in permanently well-mixed temperate coastal waters

Nitrogen uptake by net- (15–200 µm), nano- (1–15µm) and picoplankton (<1 µm) was measured overseasonal cycles at two stations with different patterns of biologicaland chemical cycles in the Morlaix Bay (western English Channel).Though assimilable dissolved N nutrient pool at both stationswas nitrate-dominated, characteristics of biomass and N uptakeby netplankton differed from conventional patterns in two respects.In the first, biomass (26–30%) and N uptake (36–43%)were less important than those of nanoplankton. In the second,the netplankton did not show any marked preference for nitrateover ammonium (nitrate to ammonium uptake ratios of 0.98 and1.08). In contrast, nanoplankton had a preference for ammoniumover nitrate (ammonium to nitrate uptake ratios of 2 and 1.2).N uptake by picoplankton was only 8% of total N uptake at bothstations and was supported mainly by regenerated N (66% ammoniumand 17% urea), with nitrate uptake detectable in only one instanceand nitrite uptake in none. Substrate-dependent uptake of ammoniumin all fractions and a higher ammonium uptake in the nanoplanktonfraction in summer at both stations when ambient ammonium concentrationswere high indicated that while nitrate may satisfy a part ofN requirements, availability of ammonium and its flux throughnanoplankton determine the magnitude of total N uptake in thesewaters. Most of the N uptake in picoplankton appears to be autotrophic,suggesting that a substantial part of heterotrophic uptake,if any, could be localized in the fractions >1 µm,and mediated by free-living and particle-bound bacteria.     

The role of phosphorus on planktonic production of the Gironde plume waters in the Bay of Biscay

More and more studies emphasize the status of phosphorus (P)as the principal limiting nutrient of phytoplankton growth,especially in coastal waters under the influence of freshwaterdischarges. The purpose of the present paper is to investigatethe role of P on planktonic production in the waters influencedby the Gironde discharges; the Gironde being one of the twolargest rivers on the French Atlantic coast. The survey is basedon several cruises made in 1998 and 1999. Two different patternswere observed for waters with salinity below and above 34.5.For waters with salinity < 34.5, P was found to be the firstlimiting nutrient of winter and spring phytoplankton blooms,based on undetectable phosphate (< 20 nM), high NO₃ : PO₄ratios, typically > 100 : 1, short phosphate turnover time(1 to 2 h), high alkaline phosphatase activities (mean of 176nM h^-1 in late May 1999) and ultimately great increases of chlorophylla (Chl a) and primary production in phosphate-enriched samplesrelative to controls. This limitation could be partly explainedby the Gironde nutrient supplies, which were phosphate deficientcompared with the mineral nitrogen(N_min : PO₄ was > 40 withina salinity range 16–33). In summer, corresponding to theperiod of low influence of Gironde supplies (low runoff anda spreading effect of the plume), phytoplankton growth wouldbe controlled by both P and nitrogen (N), according to low nitrateand the major effect of combined P+N (NH₄) enrichment on Chla and primary production compared with the addition of N orP singly. In early October, after the first autumn gales, themixed layer was enriched with a sufficient supply of nutrientsto support exponential phytoplankton growth for 4 days in enclosures.The pattern was different for waters at the limit of the Girondeplume and Atlantic oceanic waters (within salinity range 34.5–35.4).P would not be the single limiting nutrient of winter bloomsand spring phytoplankton growth since low phosphate, and alsolow nitrate and silicate, availability were recorded and phosphateaddition alone had no effect on phytoplankton biomass and productionin bioassays. The early P limitation of winter blooms had consequencesfor the phytoplankton community structure in the Gironde plumewaters (salinity < 34.5). Whereas major cells of these bloomswere greater than 20 µm in size, phytoplankton growthin spring and autumn was dominated by 3–20 µm (e.g.53% of Chl a in late April 1999) and < 3 µm cells (e.g.29% of Chl a). The decreasing size of phytoplankton cells isemphasized by the severe competition between bacteria and algaefor phosphate, since bacteria dominated phosphate uptake inspring (e.g. 87% in late April, 77% in late May). Bacteria tendedto have greater affinity for phosphate and seemed also to beP limited at certain times in spring, according to results fromphosphate enrichment bioassays in late May 1999. The alternativemethod for phytoplankton to obtain P would be the use of thedissolved organic phosphorus pool by alkaline phosphatase activity.According to the movement of ³³P after initial labelling ofmicrobial populations and a subsequent cold chase, the majortransfer of P occurred from the bacterial to the dissolved fraction.We hypothesize that algae obtain part of its dissolved organicphosphorus from bacteria-originated organic phosphorus compounds.     

Biomass estimation of components of the marine nanoplankton and picoplankton by the Utermohl settling technique

Estimation of the biomass of some components of the marine nano-and picoplankton is made for several locations using the Utermohlsettling technique. The importance of these small cells in thephytoplankton biomass is confirmed. The proportion of cellsof 2–12 µm diameter in the total phytoplankton biomassis usually lower than the proportion of primary productivityattributed to a similar size fraction in the literature. Dataare presented on geographical and vertical distributions ofsome <2 µm and 2–12 µm organisms. At theoceanic stations and at offshore sites the latter make up ahigher percentage of the total biomass than in the coastal areasbut their numerical abundance is lower. Depth distributionsare variable but oceanic profiles show a decrease in abundancewith depth. The Utermöhl technique is evaluated as a continuingmethod for biomass estimation.