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.     

Dissolved organic carbon in a humic lake: effects on bacterial production and respiration

Allochthonous matter was the main source of carbon for pelagic bacteria in a humic lake, accounting for almost 90% of the carbon required to support observed bacterial growth. The estimated contribution from zooplankton excretion was of the same magnitude as direct phytoplankton release, both accounting for 5–7% of bacterial demands for dissolved carbon. Bacteria were an important source of carbon both for heterotrophic phytoplankton and for filter feeding zooplankton species, further stressing the role of humus DOC in overall lake productivity. The high contribution of allochthonous DOC implies a stoichiometry of dissolved nutrients with a surplus of C relative to P. The high P cell quota of bacteria suggest that under such conditions they are P-limited and act like net consumers of P. Excess C will be disposed of, and bacterial respiration rate will increase following a transition from carbon-limited bacterial growth towards mineral-nutrient-limited growth. Thus the high community respiration and frequent CO₂-supersaturation in humic lakes may be caused not only by the absolute supply of organic C, but also by the stoichiometry of the dissolved nutrient pool.     

The importance of bacterial utilization of released phytoplankton photosynthate in two humic forest lakes in southern Finland

Bacterial utilization of photosynthetically fixed dissolved organic carbon (PDOC) released from natural phytoplankton assemblages was studied in two small, extremely humic, forest lakes in southern Finland. Bacterial activity (measured as uptake of ¹⁴C-glucose) and phytoplankton photosynthesis (measured as light uptake of ¹⁴CO₂) could be most effectively separated using Nuclepore filters of pore size 1–2 μm. Released PDOC was 10–67% of total phytoplankton carbon fixation during in situ experiments, and represented about 0.1% of total DOC. Net uptake of PDOC by bacteria was found to be about 20% during 24 hour laboratory incubations, although about 40% of PDOC present at the start of an experiment could be utilized by bacteria during a 24 hour period. PDOC does not provide a quantitatively important substrate supply for bacterial respiration in humic forest lakes.     

The seasonal productivity of phytoplankton in a hypertrophic, gravel-pit lake

The seasonal time course of phytoplankton primary productivitywas studied weekly in a hypertrophic, gravel-pit lake closeto Madrid, Spain. Chlorophyll a ranged 22–445 mg m^–2.Gross primary productivity attained 0.28±0.14 g C m^–2h^–1 (range: 0.06–0.60), its yearly value being 900g C m^–2, but the shallow euphotic depths and the highplankton respiration ensured that net productivity was generallylow. Respiration losses amounted to 0.31±0.24 g O₂ m^–2h^–1, with phytoplankton respiration roughly attainingone-half of overall plankton respiration. Areal phytoplanktonproductivity and plankton respiration followed a seasonal trendbut this was not the case for photosynthetic capacity. Surfacephotoinhibition was evenly distributed throughout the study.Quantum yields showed an increasing depth trend, but no seasonaltrend. Both P_max and I_k were both temperature- and irradiance-dependent.As compared with lakes of lesser trophic degree, phytoplanktonprimary production in hypertrophic lakes might be increasednot only by higher nutrient contents but also by low chlorophyll-specificattenuation coefficients and low background, non-algal attenuation,thereby allowing for higher areal chlorophyll contents and hencehigher areal productivity. Our study suggests that physical(irradiance and water column stability) as well as chemicalfeatures (dissolved inorganic carbon and soluble reactive phosphorus)may control seasonality of phytoplankton primary productionin this lake despite recent claims that only physical factorsare of significance in hypertrophic lakes. However, this doesnot explain all the variability observed and so a food web controlis also likely to be operating.     

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.     

In situ bacterial production and growth yield measured by thymidine, leucine and fractionated dark oxygen uptake

In situ bacterial net production and growth yield were measuredusing thymidine, leucine incorporation and dark oxygen consumptiontechniques in marine enclosures and in the Bay of Aarhus, Denmark.Bacterial respiration was significantly correlated with thymidine(r² = 0.42, P < 0.01, y = 0.l2x + 0.054) and leucine (r²= 0.45, P < 0.01, y = 0.09x + 0.043). The range of bacterialgrowth yield, calculated from the relationship net production/netproduction + respiration, was 0.07–0.77 with 74% of theobservations lying in the 0.15–0.45 growth yield interval.Substrate was an important determinant of growth yields. A significantdifference was found between growth yields obtained from anenclosure with added glycine (mean 0.32±0.096) and onewith added inorganic nutrients (mean 0.16±0.051) (P <0.01, t-test). Growth yield showed a weak but significant negativecorrelation with temperature (r² = 0.0.35, P < 0.001, y =–0.017 + 0.52). No correlation between chlorophyll a andgrowth yield was found (r² = 0.25, P > 0.05). The resultssuggest that thymidine and leucine techniques reflect the levelsof bacterial production to better than an order of magnitude.The variations found in the growth yield support the notionthat relying on fixed growth yields reduces the accuracy ofestimating gross bacterial production.     

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     

Pelagic carbon metabolism in a eutrophic lake during a clear-water phase

Dissolved and paniculate organic carbon (DOC and POC, respectively),primary production, bacterial production, bacterial carbon demandand community grazing were measured for 9 weeks in eutrophicFrederiksborg Slotssø. The period covered the declineof the spring bloom, a clear-water phase and a summer phasewith increasing phytoplankton biomass. The process rates andchanges in pools of organic carbon were combined in a carbonbudget for the epilimnion. The POC budget showed a close balancefor both the post-spring bloom and the clear-water phase, whilea surplus was found in the summer phase. Production of POC wasdominated by phytoplankton (2/3) compared to bacteria (1/3)during all phases, and there was a significant correlation betweenphytoplankton and bacterial production rates (r² = 0.48, P <0.039). Bacterial demand for DOC was balanced by productionand changes in the pool of DOC during the decline of the springbloom, but the calculated demand exceeded the supply by 81 and167%, respectively, during the other two periods. The discrepancywas most probably due to an underestimation of bacterial growthefficiency and an overestimation of in situ bacterial productionin carbon units. Production of bacterial substrate by zooplanktonactivity was estimated to be higher than the direct excretionof organic carbon from phytoplankton. The biological successionwas regulated by the balance between area primary productionand community grazing. The clear-water phase was initiated bya combination of low primary production due to low surface irradianceand high community grazing (100 mmol C m^–2 day^–1),which caused a decrease in phytoplankton biomass. However, dueto the high initial phytoplankton biomass, community grazingwas not high enough to cause a significant decrease in areaprimary production. The summer phase was initiated by a decreasein community grazing followed by an increase in phytoplanktonbiomass. Based on these observations and calculations of areaprimary production as a function of chlorophyll concentrations,we suggest that the possibility for zooplankton to regulatephytoplankton biomass in temperate lakes decreases with increasingnutrient level.     

Comparison of two methods to measure algel release of dissolved organic carbon and the subsequent uptake by bacteria

Two methods were used to compare phytoplankton release and bacterialuptake of extracellular organic carbon (EOC) products. A sizefractionation and an antibiotic method were used simultaneouslyduring several diel in Situ studies in Danish lakes and onecoastal area. Phytoplankton populations were very sensitiveto even small concentrations of antibiotics (1 µg ml^–1).Generally the bacterial activity was inhibited by 50%. In caseswhere the antibiotics did not affect phytoplankton photosynthesis,the two methods agreed reasonably. Bacterial respiration dataobtained with the antibiotic method were in the range of valuesfrom the literature. This supports the conclusion that the partialuncoupling of EOC release and uptake with antibiotics can beused to follow the flux of EOC in situ. The effect on the phytoplanktonmust however, be evaluated in each experiment. The methodologicalproblems of both approaches are discussed.     

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.     

Respiration and respiratory electron transport activity in marine phytoplankton: growth rate dependence and light enhancement

Respiratory electron transport system (ETS) activity and actualoxygen consumption rates were measured in batch cultures offour species of marine phytoplankton, in two different growthstages: exponential or log-phase (L) and stationary phase (S).The L cultures showed higher ETS activity and respiration ratesthan the S cultures of the same species. Among the L cultures,the higher respiration and ETS activity corresponded to thosehaving higher growth rates. The carbon-specific ETS activityand the carbon-specific respiration (h^–1) showed a cleardependence on growth rates. Samples subjected to short (10 min)exposures to high, oversaturating irradiances (1000 µEm^–2 s^–1) displayed enhanced ETS activity and respiration.The experiments show that, under the light regime at which thealgal cells grow, the respiratory ETS activity and actual oxygenconsumption in phytoplankton are strongly related to growthrate and that short, high irradiance exposures enhance boththe respiratory enzyme activity and their actual oxygen consumption.     

Picoplankton community structure along the northern Iberian continental margin in late winter-early spring

The surface distribution of autotrophic and heterotrophic picoplanktonwas assessed in 24 transects perpendicular to the coast alongthe N and NW Iberian peninsula shelf in late winter and earlyspring 2002. Community structure was analyzed by flow cytometry(FC) and found to be strongly influenced by hydrography. Typicallate winter conditions were found during the survey, characterizedby the presence of the poleward Portugal coastal counter current(PCCC) in the west and an increasing stratification eastwards.Cyanobacteria (mostly Synechococcus) dominated at low chlorophylla (Chl a) concentration whereas both the total and relativeabundance of picoeukaryotes generally increased with total phytoplanktonbiomass. Differences in the cell size of most FC-defined picoplanktonicgroups were also observed along the longitudinal and coastal–offshoregradients. The presence of Prochlorococcus (<10³ cells mL^–1)coincided with the core of the PCCC and its significant correlationwith salinity suggests its possible use as a tracer of thiscurrent. Two groups of heterotrophic bacteria were distinguishedaccording to their relative DNA content. High DNA bacteria dominatedthe community (60 ± 1% SE of total numbers), reachingmaximum values in areas under riverine influence with presumedhigher inputs of organic matter. Picoplankton biomass was dominatedby heterotrophic bacteria in the western region (58 ±3%) while autotrophic groups contributed on average 66 ±2% in the southern Bay of Biscay. The heterotrophic bacteriato phytoplankton biomass ratio decreased significantly alongthe measured range. Yet showing regional differences, the estimatedcontribution of picophytoplankton to total algal biomass washigh (mean 59 ± 4%), indicating the important role ofsmall cells at the onset of the spring bloom in these temperateshelf waters.     

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 variations of bacterial abundance and biomass and their relation to phytoplankton in the hypertrophic tropical lagoon Cienaga Grande de Santa Marta, Colombia

The seasonal development of bacteria was studied in the hypertrophiccoastal lagoon Ciénaga Grande de Santa Marta (Caribbeancoast of Colombia). This large but only 1.5 m deep lagoon issubject to strong seasonal variations of salinity from almostfully marine (April/May) to brackish conditions in October/November.Chlorophyll ranged from 6 to 182 µg L^–1, and grossprimary production amounted to 1690 g C m^–2 per year.Total bacterial number (TBN) ranged from 6.5 to 90.5 x 10⁹ cellsL^–1 and bacterial biomass (BBM) from 77 to 1542 µgC L^–1, which are among the highest ever reported for naturalcoastal waters. Neither TBN nor BBM varied significantly withsalinity, phytoplankton or seston concentrations. Only the bacterialmean cell volume showed a significant relation to salinity,being highest (0.066 µm³) during the period of increasingand lowest (0.032 µm³) during decreasing salinity. Bacterialprotein accounted for 24% (19–26%) and phytoplankton proteinfor 57% (53–71%) of total seston protein. The ratio (annualmean) of bacterial carbon to phytoplankton carbon was 0.44 (range0.04–1.43). At low phytoplankton abundance [chlorophylla (Chl a) < 25 µg L^–1], bacterial carbon wasalmost equal to phytoplankton biomass (i.e. the mean ratio was1.04). In contrast, at Chl a > 100 µg L^–1, BBMwas low compared to phytoplankton biomass (the mean ratio was0.16). In general, BBM varied less than phytoplankton biomass.Most probably, the missing correlation between bacterial andphytoplankton variables was due to (i) organic material partlyderived from allochthonous sources serving as food resourcefor bacteria and (ii) a strong resuspension of bacteria fromthe sediment caused by frequent wind-induced mixing of the veryshallow lagoon.     

Variation in temporal [14C]plankton photosynthesis among warm monomictic lakes of coastal British Columbia

Seasonal patterns of [¹⁴C]phytoplankton photosynthesis (PP)were examined in six warm monomictic lakes of coastal BritishColumbia. Four of our study lakes followed typical lake patternswith maximum PP occurring in the spring and minimal rates occurringduring the winter. However, the spring maximum occurred severalweeks earlier than lakes in other climatic regions. In addition,maximum rates of daily photosynthesis were observed to occurduring the winter months in Maxwell Lake, rather than duringthe standard growing season. All study lakes except MaxwellLake had large Daphnia in the plankton community. Maxwell wasdominated by small crustacean zooplankton implying the importanceof trophic structure in mediating seasonal patterns of productivity.The four oligotrophic lakes in our study also exhibited P-deficiencyduring winter, as indicated by P-debt bioassays and rapid ³²PO₄^3–turnover rates. Our data suggest that these coastal oligotrophiclakes were co-limited by nutrients and light during winter.The importance of winter (November–March) photosyntheticproduction to the total annual carbon budget in the six lakesstudied here is greater than that typically reported for othertemperate zone lakes. If plankton community respiration decreasesmore than photosynthetic production with wintertime temperatures,then >50% of annual net pelagic carbon fixation could occurin winter in some coastal lakes.     

Photosynthetic oxygen production and community respiration in the Indian sector of the Antarctic Ocean during austral summer

Photosynthetic oxygen production by phytoplankton and community respiration in the Indian sector of the Antarctic Ocean were estimated from changes in oxygen concentrations in light and dark bottles. Gross production varied between 0.1 and 5.1 µmol O₂ l^-1 day^-1. In the same water, community respiration (the sum of oxygen consumption by heterotrophs and phytoplankton) was 0.4-3.6 µmol O₂ l^-1 day^-1, which accounted for 47-343% of the gross production. Algal and heterotrophic respirations were distinguished using some assumptions. These estimates showed that heterotrophic respiration accounted for most of the community respiration (70-91% depending upon the assumptions), indicating that heterotrophic respiration plays an important role in the mineralization of phytoplankton production in the surveyed sea area. Gross production rate correlated with chlorophyll a concentration, showing that the photosynthetic production rate of oxygen depends on the abundance of phytoplankton. Moreover, there was a significant relationship between gross production and community respiration rates. These regression equations suggested that negative net production occurred under the usually low concentration of chlorophyll observed in the Indian sector of the Antarctic Ocean. Hence, the net exchange of carbon dioxide due to biological processes through the sea surface seemed to be not as large as expected in the Antarctic Ocean, although the number of data were limited at this stage.     

Bacterioplankton growth, grazing mortality and quantitative relationship to primary production in a humic and a clearwater lake

Bacterial growth and grazing mortality were estimated from Mayto October in two south Swedish oligotrophic lakes, one beinga clearwater lake (water colour 5–10 mg Pt l^–1 DOC2.9–3.4 mg l^–1, Secchi disk depth 5.0–9.4m) and the other a humic, brownwater lake (water colour 105–165mg Pt l^–1, DOC 13.7–22.7mg l^–1, Secchi diskdepth 1.3–2.1 m). Specific rates of growth and grazingmortality were generally similar for both lakes. However, theabundance of bacteria was consistently 2–3 times higherin the water of the humic lake, suggesting that the total productionand consumption of bacterial cells were also higher than inthe dearwater lake. The ratio of bacterial secondary productionto primary production was higher in the humic lake than in theclearwater lake, indicating that the bacterioplankton of thehumic lake utilize allochthonous substrates, in addition tosubstrates originating from autochthonous primary production.Most of the bacterial loss in both lakes could be attributedto small protozoan grazers. This implies that allochthonousand autochthonous organic carbon fixed by bacterioplankton isless important in terms of carbon flow to higher trophic levelsthan would be expected if macrozooplankton were the dominantbacterivores, providing a more direct and efficient transferof carbon to larger organisms.     

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.     

Abundance, size distribution and bacterial colonization of transparent exopolymeric particles (TEP) during spring in the Kattegat

The abundance, size distribution and bacterial colonizationof transparent exopolymeric particles (TEP) were monitored inthe Kattegat (Denmark) at weekly intervals throughout the spring(February-May) encompassing the spring diatom bloom. These recentlydiscovered particles are believed to be formed from colloidalorganic material exuded by phytoplankton and bacteria, and mayhave significant implications for pelagic flux processes. Duringthis study, the number concentration of TEP (>1 µm)ranged from 3 x 10³ to 6 x 10⁴ ml^–1 and the volume concentrationbetween 0.3 and 9.0 p.p.m.; they were most abundant in the surfacewaters subsequent to the spring phytoplankton bloom. The rangeof TEP (encased) volume concentration was similar to that ofthe phytoplankton, although at times TEP volume concentrationexceeded that of the phytoplankton by two orders of magnitude.The TEP size distribution followed a power law, with the abundanceof particles scaling with particle diameter^–(ß+1).The seasonal average estimate of ß (2.3) was not significantlydifferent from three, consistent with TEP being formed by shearcoagulation from smaller particles. However, date-specific estimatesof ß differed significantly from three, probably becauseTEP are fractal. All TEP were colonized by bacteria, and bacteriawere both attached to the surface of and embedded in TEP. Yetthe number of attached bacteria per TEP was related neitherto the surface area nor the volume, but rather scaled with TEPsize raised to an exponent of     

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.