Primary production and phytoplankton composition in relation to DOC input and bacterioplankton production in humic Lake Örträsket

1. The biomass and production of picophytoplankton, large phytoplankton and heterotrophic bacterioplankton were measured in humic Lake Örträsket, northern Sweden during four consecutive summers.
2. High flow episodes, carrying fresh dissolved organic carbon (DOC) into the lake, always stimulated heterotrophic bacterial production at the expense of primary production. Primary production never exceeded bacterial production for approximately 20 days after such an episode had replenished epilimnial DOC. We suggest that allochthonous DOC is an energy source that stimulates bacterioplankton that, because of their efficient uptake of inorganic nutrients, are then able to outcompete phytoplankton. After the exhaustion of readily available DOC, phytoplankton were able to dominate epilimnion production in Lake Örträsket.
3. Biomass production was higher when dominated by phytoplankton than by bacterioplankton, despite a similar utilization of nutrients in the epilimnion throughout the summer. We propose that different C : N : P ratios of bacterioplankton and phytoplankton permit the latter to produce more carbon (C) biomass per unit of available inorganic nutrients than bacterioplankton.     

Bacterioplankton in a small polyhumic lake with an anoxic hypolimnion

Bacterioplankton biomass and dark fixation of inorganic carbon were measured in the highly humic (water colour up to 550 mg Pt l^?1) and acidic lake, Mekkojärvi. Strong thermal and chemical stratification developed in the water column early in spring and led rapidly to anoxia in the hypolimnion, which extended to less than 1.0 m from the surface. In the epilimnion only small bacteria were abundant. In the anoxic zone both the abundance and the mean size of bacteria were considerably higher than in the epilimnion. These differences are thought to be the result of different grazing pressure from zooplankton in the two zones. In late summer a high concentration of bacteriochlorophyll d in the upper hypolimnion indicated a high density of photosynthetic bacteria. Bacterial biomass was similar to that of phytoplankton in the epilimnion, but 23 times higher in the whole water column. In August, dark fixation of inorganic radiocarbon in the anaerobic zone was 51% of the total ¹⁴C-incorporation and the contribution of light fixation was only 5.4%. In the polyhumic Mekkojarvi, bacterioplankton was evidently a potentially significant carbon source for higher trophic levels, but bacterioplankton production could not be supported by phytoplankton alone. Allochthonous inputs of dissolved organic matter probably support most of the bacterial production.     

High abundance of picoplankton-ingesting ciliates during late fall in Lake Kinneret, Israel

Small, aloricate ciliates dominated the biomass of heterotrophicprotists throughout the water column at the end of the periodof stratification in Lake Kinneret, Israel The integrated biomassof cilates was 5–20 times that of heterotrophic flagellatesDuring incubation experiments, ciliate growth rates in cpilimneticwater corresponded to population doubling times of 9.6–19.4h, while flagellate populations showed no growth. Most of thealiates were small forms (10–30 µm long), includingscuticocihates, choreotnchs, Coleps spp. and Colpoda spp., andappeared to be consuming bacteria, coccoid cyanobacteria, and<5 µm eukaryotic algae. Grazing rates of cihate assemblageson picoplankton in the epilimnion, as determined by the uptakeof fluorescently labeled bacteria and cyanobactena, ranged from62 to 86 nl cell^–1 h^–1 Colpoda steini, isolatedfrom lakewater, grew on a cultured freshwater Synechococcussp with a doubling time of 4.5 h, and a gross growth efficiencyof 48% The estimated daily requirements of ciliates for growthapproximately equalled total phytoplankton production. We calculatedthat ciliates in the epilimnion were clearing 4–10% ofthe bacterioplankton and cyanobactenal standing stocks per daySince this would not be sufficient food consumption to meetdaily carbon requirements of the aliates, it is likely thatthese organisms were also grazing a significant amount of autotrophicand heterotrophic eukaryotic cells in Lake Kinneret.     

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.     

Fish-induced changes in zooplankton grazing on phytoplankton and bacterioplankton: a long-term study in shallow hypertrophic Lake Sobygaard

The impact of fish-mediated changes on the structure and grazingof zooplankton on phytoplankton and bacterioplankton was studiedin Lake Søbygaard during the period 1984–92 bymeans of in vitro grazing experiments (¹⁴C-labelled phytoplankton,³H-labelled bacterioplankton) and model predictions. Measuredzooplankton clearance rates ranged from 0–25 ml l^–1h^–1 on phytoplankton to 0–33 ml l^–1 h^–1on bacterioplankton.The highest rates were found during thesummer when Daphnia spp. were dominant. As the phytoplanktonbiomass was substantially greater than that of bacterioplanktonthroughout the study period, ingestion of phytoplankton was26-fold greater than that of bacterioplankton. Multiple regressionanalysis of the experimental data revealed that Daphnia spp.,Bosmina longirostris and Cyclops vicinus, which were the dominantzooplankton, all contributed significantly to the variationin ingestion of phytoplankton, while only Daphnia spp. contributedsignificantly to that of bacterioplankton. Using estimated meanvalues for clearance and ingestion rates for different zooplankters,we calculated zooplankton grazing on phytoplankton and bacterioplanktonon the basis of monitoring data of lake plankton obtained duringa 9 year study period. Summer mean grazing ranged from 2 to4% of phytoplankton production and 2% of bacterioplankton productionto maxima of 53 and 88%, respectively. The grazing percentagedecreased with increasing density of planktivorous fish caughtin August each year using gill nets and shore-line electrofishing.The changes along a gradient of planktivorous fish abundanceseemed highest for bacterioplankton. Accordingly, the percentagecontribution of bacterioplankton to the total ingestion of thetwo carbon sources decreased from a summer mean value of 8%in Daphnia-dominated communities at lower fish density to 0.7–1.1%at high fish density, when cyclopoid copepods or Bosmina androtifers dominated. Likewise, the percentage of phytoplanktonproduction channelled through the bacteria varied, it beinghighest (5–8%) at high fish densities. It is argued thatthe negative impact of zooplankton grazing on bacterioplanktonin shallow lakes is highest at intermediate phosphorus levels,under which conditions Daphnia dominate the zooplankton community.     

Primary and bacterial production in lakes: are they coupled over depth?

The coupling of primary and bacterial production over depthwas examined in three lakes which differed greatly in verticalpatterns of primary productivity. We measured bacterial production,chlorophyll and light, and estimated primary production in PaulLake (Gogebic County, Michigan) and Crystal and Trout lakes(Vilas County, Wisconsin) during the summer stratification period(May–September 1991). Bacterial productivity was measuredusing the [³H]leucine incorporation method and primary productivityestimated from measured photosynthesis–irradiance relationships.Three distinct vertical patterns were observed. In Paul Lake,bacterial production was highest at the interface between theaerobic and anaerobic layers, well below the depth of maximumprimary production. In Crystal Lake, bacterial production wasuniform with depth, although primary productivity was highestin the hypolimnion. In the largest lake. Trout Lake, primaryand bacterial production tended to co-vary with maximum ratesof both processes occurring in the metalimnion. Overall, bacterialproductivity was poorly related to contemporaneous primary productionin the three lakes, suggesting that other factors, such as nutrientrecycling, phytoplankton loss rates and allocthonous loading,determine patterns in the depth distribution of bacterial productivity.     

Microplankton and primary production in the Sea of Okhotsk in summer 1994

Phytoplankton composition, density, vertical distribution andprimary production were investigated in the Sea of Okhotsk andin the adjacent northern north Pacific in July–August1994, together with measurements of density and distributionof planktonic microheterotrophs: bacteria, nanoheterotrophsand ciliates. Different phases of phytoplankton seasonal successionwere encountered during the period of investigation in variousregions of this sea. Primary production measured at 144 stationswas found to be greatest (1.5–4 g C m^-2day^-1) in areasof spring-phase succession along the Sakhalin shelf and theKashevarov bank. Periodic relapses of the spring blooms of ‘heavy’diatoms during the whole growth season were recorded over thisbank. The summer phase of the phytoplankton minimum prevailedin the central and eastern parts of the sea, manifested by thedominance of nanoflagellates in terms of phytoplankton biomass.Primary production was 0.5–1 g C m^-2 day^-1. The earlyautumn phase of succession was typical of the Kurile straitarea and the adjacent north Pacific. Primary production therevaried from 0.7 to 2 g C m^-2 day^-1. The integrated phytoplanktonbiomass in the water column varied from 9–12 g m^-2 inzones supporting the summer minimum assemblage to 15–20g m^-2 in zones of early autumn recovery of phytoplankton growth,and up to 40–70 g m^-2 in areas of remnant or relapseddiatom blooms. The numerical density of bacterioplankton wasbetween 1 x 10⁶ and 3 x 10⁶ cells ml^-1 and its wet biomass wasbetween 100 and 370 mg m^-3. In deep waters it was 8–15mg m^-3. The integrated bacterioplankton biomass in the upperwater column varied from 6 to 29 g m^-2. The numerical densityof zooflagellates varied in the upper layer between 0.8 x 10⁶and 4 x 10⁶ l^-1 and their biomass was between 20 and 50 mg m^-3.In deep waters they were still present at a density of 0.05x 10⁶ to 0.2 x 10⁶ cells l^-1. The biomass of planktonic ciliatesvaried between stations from 20 to 100 mg m^-3. The joint biomassof planktonic protozoa in the water column was between 3 and12 g m^-3 at most of the stations.     

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.     

Bacterioplankton Abundance and Activity in a Small Hypertrophic Stratified Lake

Bacterioplankton abundance and production were followed during one decade (1991–2001) in the hypertrophic and steeply stratified small Lake Verevi (Estonia). The lake is generally dimictic. However, a partly meromictic status could be formed in specific meteorological conditions as occurred in springs of 2000 and 2001. The abundance of bacteria in Lake Verevi is highly variable (0.70 to 22 × 10⁶ cells ml⁻¹) and generally the highest in anoxic hypolimnetic water. In 2000–2001, the bacterial abundance in the hypolimnion increased probably due to meromixis. During a productive season, heterotrophic bacteria were able to consume about 10–40% of primary production in the epilimnion. Our study showed that bacterioplankton in the epilimnion was top-down controlled by predators, while in metalimnion bacteria were dependent on energy and carbon sources (bottom-up regulated). Below the thermocline hypolimnetic bacteria mineralized organic matter what led to the depletion of oxygen and created anoxic hypolimnion where rich mineral nutrient and sulphide concentrations coexisted with high bacterial numbers.     

Significance of Viral Lysis and Flagellate Grazing as Factors Controlling Bacterioplankton Production in a Eutrophic Lake

The effects of viral lysis and heterotrophic nanoflagellate (HNF) grazing on bacterial mortality were estimated in a eutrophic lake (Lake Plußsee in northern Germany) which was separated by a steep temperature and oxygen gradient into a warm and oxic epilimnion and a cold and anoxic hypolimnion. Two transmission electron microscopy-based methods (whole-cell examination and thin sections) were used to determine the frequency of visibly infected cells, and a model was used to estimate bacterial mortality due to viral lysis. Examination of thin sections also showed that between 20.2 and 29.2% (average, 26.1%) of the bacterial cells were empty (ghosts) and thus could not contribute to viral production. The most important finding was that the mechanism for regulating bacterial production shifted with depth from grazing control in the epilimnion to control due to viral lysis in the hypolimnion. We estimated that in the epilimnion viral lysis accounted on average for 8.4 to 41.8% of the summed mortality (calculated by determining the sum of the mortalities due to lysis and grazing), compared to 51.3 to 91.0% of the summed mortality in the metalimninon and 88.5 to 94.2% of the summed mortality in the hypolimnion. Estimates of summed mortality values indicated that bacterial production was controlled completely or almost completely in the epilimnion (summed mortality, 66.6 to 128.5%) and the hypolimnion (summed mortality, 43.4 to 103.3%), whereas in the metalimnion viral lysis and HNF grazing were not sufficient to control bacterial production (summed mortality, 22.4 to 56.7%). The estimated contribution of organic matter released by viral lysis of cells into the pool of dissolved organic matter (DOM) was low; however, since cell lysis products are very likely labile compared to the bulk DOM, they might stimulate bacterial production. The high mortality of bacterioplankton due to viral lysis in anoxic water indicates that a significant portion of bacterial production in the metalimnion and hypolimnion is cycled in the bacterium-virus-DOM loop. This finding has major implications for the fate and cycling of organic nutrients in lakes.     

Respiratory rates in the cladoceran Ceriodaphnia dubia in Lake Rotongaio, a monomictic lake

The experimentally measured oxygen consumption rate by the cladoceran,Ceriodaphnia dubia, showed a linear increase between 5 and 20°C.Oxygen consumption rates of C. dubia were estimated in situfrom respiratory electron transport system (ETS) activity inLake Rotongaio during summer stratification and winter mixing.Oxygen consumption was 0.002 µl O₂ animal^–1 h^–1in the hypolimnion and 0.076 µl O₂ animal^–1 h^–1in the epilimnion during stratification. Implications of respiredoxygen for metabolic carbon requirements are discussed.     

Changes in bacterioplankton community structure and activity with depth in a eutrophic lake as revealed by 5S rRNA analysis

The community structure of bacterioplankton was studied at different depths (0 to 25 m) of a temperate eutrophic lake (Lake Plusssee in northern Germany) by using comparative 5S rRNA analysis. The relative amounts of taxonomic groups were estimated from 5S rRNA bands separated by high-resolution electrophoresis. Comparison of partial 5S rRNA sequences enabled detection of changes in single taxa over space and during seasons. Overall, the bacterioplankton community was dominated by 3 to 14 abundant (>4% of the total 5S rRNA) taxa. In general, the number of 5S rRNA bands (i.e., the number of bacterial taxa) decreased with depth. In the fall, when thermal stratification and chemical stratification were much more pronounced than they were in the spring, the correlation between the depth layers and the community structure was more pronounced. Therefore, in the fall each layer had its own community structure; i.e., there were different community structures in the epilimnion, the metalimnion, and the hypolimnion. Only three 5S rRNA bands were detected in the hypolimnion during the fall, and one band accounted for about 70% of the total 5S rRNA. The sequences of individual 5S rRNA bands from the spring and fall were different for all size classes analyzed except two bands, one of which was identified as Comamonas acidivorans. In the overall analysis of the depth profiles, the diversity in the epilimnion contrasted with the reduced diversity of the bacterioplankton communities in the hypolimnion, and large differences occurred in the composition of the epilimnion at different seasons except for generalists like C. acidivorans.     

Dynamics of inorganic phosphorus in pelagic communities of the Sea of Okhotsk

Inorganic phosphorus uptake and regeneration in the OkhotskSea waters were investigated in July–August 1994 withthe use of radioisotopic techniques. The rates of PO₄-P uptakeby microplankton in the upper mixed layer were between 1.5 and6.6 µg P l^-1 day^-1 (average 2.75) in areas of diatom dominance,and between 0.68 and 1.68 µg P l^-1 day^-1 (average 1.16)in areas of intense warming and summer phytoplankton minimum.The residence time of PO₄-P standing stock in water at differentstations varied between 1.5 and 24 days (mean 9 days). The shareof bacterioplankton contributing to total PO₄-P uptake was 50%in areas of the summer phytoplankton minimum and 20–30%in areas of diatom dominance. The PO₄-P regeneration rate wasmeasured first time experimentally in the temperate sea. Itsrates varied from 0.30 to 1.65 µg P l^-1 day^-1. In areasof diatom dominance, it compensated with 30–60% of PO₄-Puptake. In zones of summer phytoplankton minimum and in thelayers of deep chlorophyll maxima at 10–25 m depths, thePO₄-P regeneration rate often exceeded its uptake. Primary phytoplanktonproduction correlated well with PO₄-P uptake values in the uppermixed layer, while no correlation was found between primaryproduction and the ambient PO₄-P content in water.     

Seasonal fluctuations in the biomass and metabolic activity of bacterioplankton and phytoplankton in a well-mixed estuary: the Ems-Dollard (Wadden Sea)

The biomass and metabolic activity of bacterioplankton weremeasured over 1 year in the Ems Dollard Estuary, a part of theDutch-German Wadden Sea. Very productive phyzoplankton blooms,composed mainly of diatoms and the haptophycean alga Phaeocystispouchetii, are a feature of the estuarine section studied. Theproduction of bacterial populations, as measured using the [³H]thymidinemethod broadly followed the phytoplankton density during bloomsin spring and late summer. There was no indication of a disproportionateincrease in bacterial production during the bloom or declineof particular algal species. The rate at which ¹⁴C-labelledglucose, glutamate and leucine were incorporated by bacterialpopulations, measured as a metabolic potential, varied seasonally,but did not precisely follow the rate of [³H]thymidine incorporation.The utilization of the absorbed ¹⁴C substrates for respirationor for cellular synthesis was constant over a prolonged periodof bacterial growth; apparent yields of 0.7, 0.4 and 0.8 weremeasured for glucose, glutamate and leucine, respectively. Inthe colder season most of the absorbed substrate was respired.The production by pelagic bacteria was 60 gC m^–2 y^–1,a value that amounted to 12% of the pelagic primary production.A preliminary experiment with a mixed culture of the abundantlyoccurring diatom species Thalassiosira excentrica and a marinespirillum, indicated that part of the algal exudates were rapidlyconverted by the bacteria, but another part resisted degradation.The incomplete bacterial degradation of algal exudates togetherwith the short residence time of the estuarine phytoplanktonmay contribute to the apparently incomplete mineralisation ofthe primary production in the water.     

Dynamics of autotrophic picoplankton in Lake Constance

The vertical distribution, biomass concentrations and growthrates of autotrophic picoplankton (APP) were investigated duringthe growing season (March-October) in Lake Constance in differentdepths. Cell numbers determined by epifluorescence microscopyvaried between 1.0 x 10³ and 1.6 times; 10⁵ cells ml^–1depending on season and water depth. Highest concentrationswere recorded above the thermodine in late summer. Numerically,APP consisted almost exclusively of chroococcoid cyanobactena.During lake stratification several peaks of biomass concentrationsoccurred in epilimsietic waters at intervals of 6–8 weeks.In-situ experiments using a dilution technique and dialysisbags revealed that during summer APP population dynamics wereprimarily driven by combined changes of their growth and grazingrates, whereas temperature was less important. Gross growthrates varied between 0.006 and 0.051 h^–1, grazing ratesbetween 0.002 and 0.053 h^–1. On average APP productionwas completely removed by grazing within the microbial community.Ciliates, heterotrophic nanoflagellates and rotifers have beenidentified as the major consumers of APP cells. APP biomassis small compared to larger phytoplankton, ranging from ito5% of total phytoplankton biovolume. Due to its high gross growthrates, which are on the same level as those of free-living pelagicbacteria, APP contributes slightly more to overall primary productionwith maximum percentages of {small tilde}15% in late summer.     

Regeneration and uptake of ammonium by plankton in an Amazon floodplain lake

Planktonic regeneration and uptake of ammonium was measuredin Lake Calado on the floodplain of the Amazon River (Brazil).In the epilimnion of the lake, regeneration and uptake averaged0.86 ± 0.15 and 1.9 ± 0.4 µmol NH₄1^–1h^–1 respectively. Uptake exceeded regeneration under theammonium-enriched conditions of our experimental incubations,but uptake and regeneration are in near-balance at in situ ammoniumconcentrations of <1 µM. In experiments that simulatedlake overturn, samples from the meta- and hypolimnion had higherrates of regeneration (1.4 ± 0.6) and lower rates ofuptake (0.89 ± 0.53) than epilimnetic samples. Duringinflow of water from the Amazon River and a local stream, ammoniumregeneration rates were similar to those measured in the epilimnion(0.84 ± 0.22), but the uptake rates were lower (0.49± 0.13). Although regeneration always exceeded uptakein the overturn and inflow experiments, ammonium did not accumulate.Rapid isotope dilution of the ammonium pool occurred in theincubations, and the effect of not accounting for isotope dilutioncould result in underestimating uptake rates by a factor of2–5 in the epilimnion of this tropical lake.     

Photosynthetic algal production, accumulation and release of phytoplankton storage carbohydrates and bacterial production in a gradient in daily nutrient supply

In a mesocosm experiment providing a gradient of semi-continuousaddition of mineral nutrient, production rates and mortalityof phytoplankton were estimated. Heterotrophic bacterial biomassand production rates and their responses to the mineral nutrientsadditions were also estimated. The purpose of the experimentwas to establish responses of the major biological factors asa function of nutrient amendments. Initial primary productionwas 0.47 µg C L^–1 day^–1. In the most fertilizedmesocosm, phytoplankton biomass increased at a specific rateof 0.4 day^–1 during the first week of the experiment,and on day 9 primary production reached a peak at 1027 µgC L^–1 day^–1. The responses in the other fertilizedmesocosms were intermediate, and in an unfertilized controlthe variables measured stayed almost constant throughout theexperiment. The termination of the blooms in the fertilizedmesocosms was a consequence of nitrogen limitation, and nitrogenlimitation subsequently induced storage of intracellular organicmaterial in the phytoplankton. In the mesocosm receiving thehighest daily dose of nutrients, strong post-bloom nutrientlimitation resulted in high phytoplankton mortality, and releaseof organic material from the algae supported the gradient’shighest heterotrophic bacterial production.     

Copepod abundance in the western Irish Sea: relationship to physical regime, phytoplankton production and standing stock

The distinct patterns of stratification in the North Channeland stratified region of the western Irish Sea influence theseasonal abundance of phytoplankton. The 3–4 month productionseason in the stratified region was characterized by productionand biomass peaks in the spring (up to 2378 mg C m² day^–1and 178.4 mg chlorophyll m^–2) and autumn (up to 1280 mgC m^–2 day^–1 and 101.9 mg chlorophyll m^–2).Phytoplankton in the North Channel exhibited a short, late productionseason with a single summer (June/July) peak in production (4483mg Cm^–2 day^–1) and biomass (–160.6 mg chlorophyllm^–2). These differences have little influence on copepoddynamics. Both regions supported recurrent annual cycles ofcopepod abundance with similar seasonal maxima (182.8–241.810³ind. m^–2) and dominant species (Pseudocalanus elongatusand Acartia clausi). Specific rates of population increase inthe spring were 0.071 and 0.048 day¹ for the North Channel andstratified region, respectively. Increased copepod abundancein the stratified region coincided with the spring bloom, andwas significantly correlated with chlorophyll standing stock.Increased copepod abundance preceded the summer production peakin the North Channel. This increase was not correlated withchlorophyll standing crop, suggesting that a food resource otherthan phytoplankton may be responsible for the onset of copepodproduction prior to the spring bloom. Hetero-trophic microplanktonas an alternative food source, and advection of copepods fromthe stratified region, are proposed as possible explanationsfor copepod abundance increasing in advance of the summer peakin primary production.     

Occurrence of mixotrophic flagellates in relation to bacterioplankton production, light regime and availability of inorganic nutrients in unproductive lakes with differing humic contents

1. Field data from five unproductive Swedish lakes were used to investigate the occurrence of mixotrophic flagellates in relation to bacterioplankton, autotrophic phytoplankton, heterotrophic flagellates and abiotic environmental factors. Three different sources of data were used: (i) a 3‐year study (1995–97) of the humic Lake Örträsket, (ii) seasonal measurements from five lakes with widely varying dissolved organic carbon (DOC) concentrations, and (iii) whole lake enrichment experiments with inorganic nutrients and organic carbon. 2. Mixotrophic flagellates usually dominated over autotrophic phytoplankton in Lake Örträsket in early summer, when both bacterial production and light levels were high. Comparative data from the five lakes demonstrated that the ratio between the biomasses of mixotrophic flagellates and autotrophic phytoplankton (the M/A‐ratio) was positively correlated to bacterioplankton production, but not to the light regime. Whole lake carbon addition (white sugar) increased bacterial biomass, and production, reduced the biomass of autotrophs by a factor of 16, and increased the M/A‐ratio from 0.03 to 3.4. Collectively, the results indicate that the dominance of mixotrophs among phytoplankton was positively related to bacterioplankton production. 3. Whole lake fertilisation with nitrogen (N) and phosphorus (P) demonstrated that the obligate autotrophic phytoplankton was limited by N. N‐addition increased the biomass of the autotrophic phytoplankton but had no effect on mixotrophic flagellates or bacteria, and the M/A‐ratio decreased from 1.2 to 0.6 after N‐enrichment. Therefore, we suggest that bacteria under natural conditions, by utilising allochthonous DOC as an energy and carbon source, are able to outcompete autotrophs for available inorganic nutrients. Consequently, mixotrophic flagellates can become the dominant phytoplankters when phagotrophy permits them to use nutrients stored in bacterial biomass. 4. In Lake Örträsket, the biomass of mixotrophs was usually higher than the biomass of heterotrophs during the summer. This dominance could not be explained by higher grazing rates among the mixotrophs. Instead, ratios between mixotrophic and heterotrophic biomass (the M/H‐ratio) were positively related to light availability. Therefore, we suggest that photosynthesis can enable mixotrophic flagellates to outcompete heterotrophic flagellates.