Grazing on bacteria by flagellates and cladocerans in lakes of contrasting food-web structure

We tested the hypothesis that grazing on bacteria would varybetween lakes with differing plankton community structures.Paul and Tuesday lakes (Gogebic County, MI) are respectivelydominated by piscivorous and planktivorous fish. Consequently,zooplankton in Paul are primarily large daphnids, while zooplanktonin Tuesday are primarily small cladocerans and copepods. Wemeasured flagellate grazing on bacteria using a fluorescentminicell method, while cladoceran grazing was estimated fromthe relationship between body length and filtering rate. Wepredicted that cladoceran grazing on bacteria would be higherin Paul, and flagellate grazing would be higher in Tuesday.Cladoceran grazing on bacteria was important in both lakes contraryto our initial expectation. Large populations of the small cladoceran,Bosmina longirostris, in Tuesday exerted a grazing pressure(0.18–35x10⁶ bacteria 1^–1 h^–1) approximatelyequal to that of the large cladoceran, Daphnia pulex, in Paul(0.34–30x10⁶ bacteria 1^–1 h^–1). Flagellategrazing was higher in Tuesday as predicted (range: Paul, 0.1–6x10⁶bacteria 1^–1 h^–1; Tuesday, 0.2–20x10⁶ bacteria1^–1 h^–1). However, there was not a simple relationshipbetween total abundance of flagellates and total grazing rates.High community grazing by flagellates occurred when attachedchoanoflagellates were present. These flagellates had higheringestion rates than free forms. We find no clear evidence thatdifferences in food-web structure between the two lakes influencethe process of grazing on bacteria. Instead, our results emphasizethe significance of cladocerans and attached flagellates asconsumers of bacteria in freshwater ecosystems.     

Indirectly fluorescently labelled flagellates (IFLF): a tool to estimate the predation on free-living heterotrophic flagellates

A procedure was developed to estimate the direct grazing impacton free-living heterotrophic nanoflagellates (HNF). Culturedflagellates were labelled by feeding on brightly fluorescingbacteria (FLB) and then offered as indirectly fluorescentlylabelled flagellates (IFLF) to potential predators of HNF. Thenumber of FLB in the predators' food vacuoles could be convertedinto IFLF uptake and consumption of HNF. This new techniquewas used to study the HNF-ciliate relationship in the pelagiczone of Lake Constance. Three groups of ciliates were detectedas HNF grazers: small representatives of the genus Strobilidium.a small Haltena-like ciliate (probably Halteria grandinella)and a Codonella sp. Tintinnidium sp. group The ingestion ofHNF by these groups of ciliates ranged between 3 and 15, 3 and39, and 3 and 7 HNF ciliate^–1 h^–1; respectively.The IFLF method allows the direct determination of ingestedflagellate prey in the food vacuoles of their predators. Becauseindigenous living prey organisms were used, tracer discriminationcan be reduced.     

Pigmented nanoflagellates in the coastal western subtropical Pacific are important grazers on Synechococcus populations

Although the key grazers on Synechococcus and other planktonicmarine bacteria are generally thought to be nanoflagellates(both non-pigmented and pigmented) as well as ciliates, we previouslyfound in our western subtropical Pacific coastal study sitethat ciliates exerted almost no grazing pressure. In this study,we used fluorescently labeled particles (FLP) as Synechococcus-sizedmimics to examine the contribution of pigmented (may includeautotrophic and mixotrophic spp.) versus non-pigmented (heterotrophic)nanoflagellate grazing to Synechococcus morality. During thewarm season from June to September, > 50% of the nanoflagellatepopulation was pigmented (1.8–2 x 10³ versus 1.2–1.6x 10³ cells mL^–1). Consumption, or clearance rates perpigmented nanoflagellate, varied considerably (0.50–46.90nL cell^–1 h^–1), with the highest rates in June.Raw data showed pigmented nanoflagellate grazing to accountfor 2–94% (mean 43%) of Synechococcus production fromMay to October. Pigmented nanoflagellates consumed 12.5-foldmore Synechococcus than did ciliates. This study provides thefirst evidence that pigmented nanoflagellates are key grazersof Synechococcus populations in subtropical western Pacificcoastal waters.     

Growth and grazing rates of Protoperidinium hirobis Abe, a thecate heterotrophic dinoflagellate

Growth and feeding rates of a laboratory-reared small thecateheterotrophic dinoflagellate, Protoperidinium hirobis Abè,grown on the diatom Leptocylindrus danicus, were measured inbatch cultures. Ingestion rates were determined directly bythe enumeration of empty diatom frustules produced by dinoflagellatefeeding. Both growth and feeding rates saturated at diatom concentrationsof {small tilde} 10⁴ cells ml^–1, and reached maximum valuesof 1.7 divisions day^–1 and 23 diatoms grazer^–1 day^–1,respectively. This rate of cell division is notably high comparedto photosynthetic dinoflagellates, which seldom grow fasterthan 1 division day^–1. A maximal clearance rate of 0.5µl h^–1 was measured. Mean cell size varied proportionallywith food abundance, with food-saturated cells having doublethe mean volume of food-depleted cells. Tuning of cell divisionand grazing rate patterns were also examined; while mitosisoccurred chiefly during the dark period, no diel variationsin feeding rate were detected. These rates represent the firstdirect growth and ingestion measurements to be made for a thecateheterotrophic dinoflagellate. They serve to underscore one functionthese dinoflagellates perform within the microzooplanktonicfood web: that of transforming large diatoms into particlesmore easily ingested by microzooplankters.     

Nanoheterotroph grazing on bacteria and cyanobacteria in oxic and suboxic waters in coastal upwelling areas off northern Chile

The vertical distribution and abundance of microbial assemblagesand the grazing of nanoheterotrophs upon prokaryotes in oxicand suboxic waters were examined in two coastal upwelling areasoff northern Chile where a shallow Oxygen Minimum Zone (OMZ)is characteristic. Prokaryotic prey included bacterioplanktonand cyanobacteria (Synechococcus); both displayed a bimodaldistribution, with abundance maxima above and within the upperOMZ. Flagellates numerically dominated the nanoplankton andwere mostly concentrated in the oxic layer. Mean ingestion ratesof cyanobacteria by nanoflagellates (vacuole content method)ranged from 0.2 to 1.1 cells flagellate^–1 h^–1 andmean consumption rates (34–160 cells mL^–1 h^–1)were four times higher in the oxic layer. With the selectiveinhibitors technique, specific grazing rates on bacteria werelow (<0.1 h^–1) and consumption did not control bacterialproduction in the surface layer but did so in the suboxic layer(accounting for >100% of bacterial production). With thesame method, the specific grazing rate on cyanobacteria rangedbetween zero and 0.23 h^–1 with no clear differences betweenoxygen conditions; prey growth and production were always higherthan the grazing pressure (accounting for <17% of cyanobacterialproduction). The impact of grazing by nanoheterotrophs in regulatingthe production of prokaryotes in oxic and suboxic waters inthis region is discussed.     

Abundance and productivity of heterotrophic nanoplankton in Georgia coastal waters

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

Microbial dynamics during the summer ice-loss phase in maritime Antarctic lakes

The dynamics of bacterioplankton and protozooplankton in twomaritime Antarctic lakes (Heywood Lake and Sombre Lake, SignyIsland, South Orkneys) were studied during the phase of icebreak-out (December and early January 1994/95). The lakes aresuffering animal-induced (fur seal) eutrophication, though HeywoodLake is most severely affected. Both lakes had morphologicallydiverse bacterial communities which increased during the studyperiod, reaching maxima of 80 x 10⁸ l^–1 in Heywood Lakeand 31.8 x 10⁸ l^–1 in Sombre Lake. Heterotrophic nanoflagellates(HNAN) reached a peak in late December with maxima of 40.6 x10⁸ l^–1 in Sombre Lake and 174 x 10⁵ l^–1 in HeywoodLake. Phototrophic nanoflagellates (PNAN) peaked in late Decemberafter ice loss in Heywood Lake (63 x 10⁵ l^–1), which coincidedwith a peak in a bloom of Chroomonas acuta which reached abundancesof 1.0 x 10⁸ l^–1. In Sombre Lake, ice persisted for alonger period and here PNAN reached their highest density atthe end of the study period (around 70.0 x 10⁵ l^–1). Ciliateabundance reached high levels in Heywood Lake (>60001^–1),while in Sombre Lake maximum abundance was 568l^–1. Protozooplanktondiversity was greater in the less-enriched Sombre Lake. Grazingrates of ciliates averaged 70.6 bacteria indiv.^–1 h^–1in Heywood Lake and 119.3 bacteria indiv.^–1 h^–1in Sombre Lake. The difference was a reflection of the differenttaxonomic make-up of the community in the lakes. HNAN grazingrates varied between 0.51 and 0.83 bacteria indiv.^–1 h^–1in Sombre and Heywood Lakes, respectively. Specific growth rates(r) h^–1 in Sombre Lake were 0.028 for ciliates and 0.013for HNAN, and in Heywood Lake 0.010 for ciliates and HNAN 0.012.These growth rates result in doubling times ranging between38 and 69 h for ciliates and around 55 h for HNAN.HNAN grazingon bacteria was curtailed in Heywood Lake in early January asa result of predation by microcrustacean larvae feeding on theplankton. Thus, for a short phase top-down control was apparentin the dynamics of Heywood Lake, a feature uncommon in Antarcticlake ecosystems. The impact of natural eutrophication on thesesystems is discussed in relation to other unaffected Antarcticlakes.     

Factors affecting the bacteria-heterotrophic nanoflagellate relationship in oligo-mesotrophic lakes

The coupling between bacteria and heterotrophic nanoflagellates(HNF) was examined in nine lakes of low productivity for evidenceof the effects of various metazooplankton (i.e. rotifers, cladoceransand copepods) on this relationship. We considered the size ofcladocerans and, in contrast to most previous across-systemstudies, the three strata of the water column (i.e. epilimnion,metalimnion and hypolimnion). Rotifers were numerically dominantin all lakes and accounted for 45–84% of total metazooplanktonabundance, while the abundance of large cladocerans was relativelylow, ranging from 0.066 to 15.2 ind. L^–1. The across-lakerelationship between bacteria and HNF was significant in thedeeper strata (meta- and hypolimnion) but not in the epilimnionand in the two groups of lakes separated on the basis of theiraverage number of large cladocerans (<5 and >5 ind. L^–1,respectively). The results confirmed the negative impacts oflarge cladocerans on HNF, but also showed that rotifers, probablythrough grazing on HNF, may be an important factor causing variationin the bacteria–HNF relationship in unproductive waters.Quadratic models best described the relationships between metazooplanktonand the ratio of bacteria to HNF. This ratio seemed to be aresult of complex interactions between several factors, includingthe zooplankton composition and abundance and the depth of thelake. Indeed, this ratio significantly decreased across lakes,with increase in depth. In addition, shallower lakes (having<5 large cladocerans L^–1 and fewer Polyarthra vulgaris)tended to have more bacteria and HNF and a higher ratio of bacteriato HNF than deeper lakes (which had >5 large cladoceransL^–1 and substantial proportions of P. vulgaris). We suggestthat the epilimnion, metalimnion and hypolimnion of lakes betaken into account when analysing the bacteria–HNF relationshipas well as the cascading effects of zooplankton on microbialcommunities.     

Short-term changes of protozoan control on autotrophic picoplankton in an oligo-mesotrophic lake

In May 1994, we investigated the short-term development of theplanktonic community in the epi- and metalimnion of an oligo-mesotrophiclake (Piburger See, Tyrol), focusing on trophic links betweenprotists and picoplankton, but also including phyto- and zooplankton.Uptake experiments with fluorescently labelled bacteria (FLB)and picocyanobacteria (FIC) were performed in order to comparethe importance of both prey types as carbon sources for bacterivorousprotists. Heterotrophic nanoflagellates (HNF) were responsiblefor {small tilde}90% of total protozoan picoplanktivory (FLB+ FLC); ciliates accounted for {small tilde}10%. A selectivityindex related to prey density showed that both HNF and ciliatesclearly preferred FLC over FLB. The mean cell size of autotrophic(prokaryotic) picoplankton (APP) was nearly three times larger(0.323 µm³) and much less variable than mean bacterialcell volume (0.122 µm³). Although APP biomass was on averageonly 8.6% of total picoplankton biomass, pico-cyanobacteriaaccounted for a mean 15.9% of total HNF carbon uptake. We calculatedthat total HNF grazing could match potential APP maximum growthrates at the beginning of the study period. A strong decreasein HNF individual clearance rate (CR) on APP was clearly relatedto a fall in the percentage of choanofiagellates (from 75 to{small tilde}10% of the HNF community). A simultaneous decreasein HNF biomass and CR was followed by a steep increase in APPabundance; APP abundance and HNF biomass were highly negativelycorrelated both in the epi- and the metalimnion (r_{1 EM} = –0.879,r_{1 META} = =0.907; P = 0.001). Total rotifer abundance increasedby a factor of 50 within 2 weeks and was also negatively correlatedwith HNF biomass (r_{1 EM} = –0.852, P < 0.001; r_{1 META}= –0.659, P < 0.05). HNF grazing was found to exerta strong short-term control on picocyanobacteria and this linkwas probably broken by an increase in metazooplankton, especiallydue to rotifer predation on HNF.     

Pyrosoma atlanticum (Tunicata, Thaliacea): grazing impact on phytoplankton standing stock and role in organic carbon flux

Pyrosomas are the large group of pelagic tunicates whose trophicrole in pelagic communities has not yet been sufficiently studied.We ran across a local area of high concentration of the mostwidespread and commonest species of pyrosomas, Pyrosoma atlanticum,450 miles off the Congo river mouth. The following was estimated:gut pigment content, defecation rate, organic carbon and pigmentcontent of fecal pellets, and sinking rate. Based on these dataand the measured number of pyrosomas colonies the grazing impacton phytoplankton and the fecal pellet flux were calculated.During the night swarms of 50–65 mm P.atlanticum removed53% of phytoplankton standing stock in the 0–10 m layer;sparsely distributed pyrosomas consumed only 4%. The grazingimpact in the 0–50 m layer was only 12.5 and <1% respectively.The fecal pellet flux resulting from nocturnal feeding of P.atlanticumwhile swarming made up 1.4–1.6 x 10⁶ pellets m^–210 h^–1 or 305–1035 mg C m^–2 10 h^–1 and1.4 x 10⁵ pellets m^–2 10 h^–1 or 87.4 mg C m^–210 h^–1 while non-swarming. Incubation experiments showedthe rapid degradation of fecal pellets at 23°C: the lossof pigment and carbon content was {small tilde}60–70%after 45 h. We believe that given the sinking rate of 70 m day^–1the main part of fecal material does not leave the upper watercolumn and is retained in the trophic web of the epipelagiclayer.     

Laboratory-measured grazing and ingestion rates of the salp, Thalia democratica Forskal, and the doliolid, Dolioletta gegenbauri Uljanin (Tunicata, Thaliacea)

Grazing and ingestion rates of laboratory-born Thalia democraticaaggregates and Dolioletta gegenbauri gonozooids, phorozooidsand oozooids were determined while fed Isochrysis galbana (4–5µm diameter) alone or in combination with Peridinium trochoideum(16–18 µm diameter) at concentrations of 0.15–0.70mm³ x 1^–1. Grazing rates (ml x zooid^–1 x 24 h ^–1)ranged from 10 to 355, and at zooid weights greater than 5 µgcarbon were in order oozooid > gonozooid > aggregate.Grazing rates increased exponentially with increasing zooidweight. Weight-specific grazing rates (ml x µgC^–1x 24 h^–1) were independent of the four-fold initial foodconcentration. Mean weight-specific grazing rates increasedlinearly with increasing zooid weight for the aggregates andoozooids, but gonozooid mean rates were independent of zooidweight. Aggregate and gonozooid ingestion rates (10⁶ µm³x zooid^–1 x 24 h^–1) ranged from 4 to 134 while oozooidrates ranged from 3 to 67. All ingestion rates were independentof the initial food concentration but increased linearly withincreasing zooid weight at similar rates. All mean weight-specificingestion rates (ml x µgC^–1 x 24 h^–1) wereindependent of zooid weight. The mean aggregate daily ration(µgC ingested x µg body C^–1) was 59% and themean doliolid ration was 132%. Field studies indicate that normalconcentrations of D. gegenbauri in the Georgia Bight clear theirresident water volume (1 m³) in about 4 months, but that highlyconcentrated, swarm populations which occur along thermohalinefronts clear their resident water volume in less than 1 day. ¹Current address: MacLaren Plansearch Ltd., P.O.Box 13250, sta.A.,St.John's, Nfld. A1B 4A5     

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.     

Density and distribution of bacterioplankton and planktonic ciliates in the Bering Sea and North Pacific

The total number of planktonic bacteria in the upper mixed layerof the Bering Sea during the late spring-early summer periodranged between 1 and {small tilde}4 x 10⁶ ml^–1 (biomass10–40mg C m^–3). In the northern Pacific, along 47–52⁶N,the corresponding characteristics of the bacterioplankton densityin the upper mixed water layer were: total number 1–2x 10⁶ cells ml^–1 and biomass 15–46mg C m^–3Below the thermocline at 50–100 m, the density of bacterioplanktonrapidly decreased. At 300 m depth, it stabilized at 0.1–0.2x 106 cells ml^–1. The integrated biomass of bacterioplanktonin the open Bering Sea ranged between 1.2 and 3.6 g C m^–2(wet biomass 6–18 g m^–2) Its production per dayvaried from 2 to 23 mg C m^–3 days^–1 in the upper0–100 m. The numerical abundance of planktonic ciliatesin this layer was estimated to be from 3 to l0 x 10³ cells l^–1,and in the northern Pacific from 0.4 to 4.5 x 10³ l^–2.Their populations were dominated by naked forms of Strombidium,Strombilidium and Tontonia. In some shelf areas, up to 40% ofthe total ciliate population was represented by the symbioticciliate Mesodinium rubrum. The data on the integrated biomassof basic groups of planktonic microheterotrophs are also presented,and their importance in the trophic relationships in pelagiccommunities of subarctic seas is discussed.     

Food selection and growth of the planktonic ciliate Coleps hirtus isolated from a monomictic subtropical lake

A strain of Coleps hirtus (Ciliophora, Prorodontida) was isolatedfrom the epilimnion of monomictic Lake Kinneret. Growth of thisciliate was tested in response to 12 species of planktonic algaeand seven species of cultured bacteria from lake isolates whichwere offered as food. Eight species of algae (one Cryptophyceaeand seven Chlorophyceae) and four bacteria supported good toexcellent growth of C.hirtus. Growth rates (µ) and doublingtimes (DT) ranged from 0.008 to 0.029 h^–1 and from 23.9to 90.8 h respectively. C.hirtus was able to grow on bacteriaat concentration levels as low as 2–8 x 10⁵ cells ml^–1.No correlation was observed between growth rate of C.hirtusand cell volume of the prey. ^aPresent address: Istituto di Ecologia, Universita di Parma,43100 Parma, Italy     

Bacterial growth and losses due to bacterivory in a mesotrophic lake

Bacterial secondary production and rates of bacterivory weredetermined from samples collected from mesotrophic Lake Arlington.Bacterial production and losses were determined by comparingthe growth of natural bacterial assemblages in the presenceof predators (unfiltered samples) to growth in the absence ofpredators (water filtered through 1.0 (im porosity filters).Growth rates of heterotrophic nanoflagellates (HNF) were estimatedfrom growth in the absence of predators (water filtered through5.0 µm porojity filters). Bacterial growth rates rangedbetween 0.002 and 0.069 h^–1 and averaged 0.026 h^–1.HNF grew at rates ranging between 0.003 and 0.107 h^–1and averaged 0.028 h^– Grazing rates ranged between 0.002and 0.043 h^–1, and averaged 0.018 h. The annual averagerate of bacterial biomass synthesis was 3.2 –g Cl^–h^–1 and {small tilde}69% of this production was grazed.Temporal changes in growth and grazing rates suggest a tightlycoupled predator-prey linkage in this lake. ¹Present address: Hydrobiological Institute, Czech Academy ofSciences, Na sddkach 7, 370 05 teski Budjovice, Czech Republic     

Growth and production of heterotrophic nanoflagellates in a meso-eutrophic lake

The growth of heterotrophic nanoflagellates (HNF) in mesotrophicLake Constance was measured in situ during a 13 month period.Experiments were conducted with 10 µm pre-filtered lakewater incubated in diffusion chambers at 3 m water depth atthe sampling location for 24 h. Growth rates were calculatedfrom changes in cell numbers occurring during the period ofincubation. Growth rates of all dominant taxa showed pronouncedseasonal variation (–0.13 to 1.76 day^–1 and weregenerally highest in summer at high water temperatures. In situgrowth rates were well below maximum growth rates known forthe respective and similar species from laboratory experiments.While water temperature was a key parameter positively relatedto the growth of all HNF species, the effect of various potentialfood items was taxon specific and less clear. Bacterial abundancewas equally important as temperature for growth in the smallbactenvorous Spumella sp., but was insignificant for growthrates of the larger omnivorous Kathablepharis sp. In Spuniellasp., 84% of the observed seasonal variation of its growth ratecould be explained by temperature and bacterial food supply.Based on these results, a multiple linear regression equationwith temperature and bacterial concentration as dependent variableswas calculated for the growth rate of Spumella. Taxon-specificproduction rates were derived from growth rates and averagebiomass of these two species, and compared to total HNF productionestimated from previously measured community growth rates andbiomass in Lake Constance. Production peaks of Spumella sp.and Kathablepharis sp. alternated seasonally. Total HINF productionranged from –0.01 to 10 mg C m^–3 day^–1. Theaverage seasonal production varied between 1.4 and 33 mg C m^–3day^–1 over 6 consecutive years. These small protozoa thuscontribute a substantial amount to total zooplankton productionin Lake Constance.     

Annual variation in the abundance and size of heterotrophic nanoflagellates on the SW coast of Finland, the Baltic Sea

Annual variation and vertical distribution in the abundanceand cell volume of heterotrophic nanoflagellates (HNF) was studiedon the SW coast of Finland, the Baltic Sea. HNF cell numbersand mean cell volume varied annually from 90 to 10⁴ cells ml^–1,and from 3 to 32 µm³, respectively, with maxima in earlysummer. The proportion of choanoflagellates in the HNF communitywas 0–23%. Statistical analysis revealed the verticaldifferences in HNF abundance to be insignificant, but verticaldifferences in the size structure of HNF communities were found,especially during thermal stratification. The majority (>80%)of HNF were small (maximum dimension 2–4 µm); theproportion of large (>7 µm) cells were only 2–4%of the HNF abundance. An empirical equation for the relationshipbetween HNF cell length and volume is presented, and the measurementof flagellate volume by epifluorescence microscopy is discussed.     

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.     

Food size selectivity of Artemia franciscana at three developmental stages

Food size selectivity was examined in Artemia franciscana metanaupliiat three different developmental stages. Clearance rates weredetermined in short-term experiments either by measuring thedecrease in concentration of live particles and plastic beads,or by measuring the radioactivity accumulated in animals thatgrazed ¹⁴C-labelled live particles. The maximum clearance rateofA.franciscana metanauplii increased during development andwas measured at 50–63 µl ind.^–1 h^–1,254 µl ind.^–1 h^–1 and 1.48–2.10 ml ind.^–1h^–1 in 2-, 4- and 7-day-old metanauplii, respectively.A preference for particles with a diameter of 4–8 µmwas observed at all three developmental stages. The abilityof A.franciscana metanauplii to graze bacterial particles wasalso demonstrated, although the efficiency in grazing such smallparticles was low compared to microalgae (28, 20 and 9% of themaximum clearance rate in 2-, 4- and 7-day-old metanauplii,respectively). Electron microscopy showed that the inter-setulardistance in antennae and thoracopods was 0.20 ± 0.07,0.16 ± 0.05 and 0.18 ± 0.04 µm in 2-, 4-and 7-day-old metanauplii, respectively, and accordingly independentof stage.     

The Effects of Light Intensity and External Potassium Level on Root/Shoot Ratio and Rates of Potassium Uptake in Perennial Ryegrass (Lolium perenne L.)

Loliun perenne L. (cv.S. 23) was grown on vermiculite in winterin a heated greenhouse for 8 weeks under factorial combinationsof two potassium regimes (nominally 6 parts/10⁶ and 156 parts/10⁶in Hewitt's solution) and three densities of artificially supplementedvisible radiation flux (36.1, 7.3, and 2.2 W m^–2). Growthand potassium uptake were studied through the calculation ofvarious growth functions from fitted curves. There was little effect of potassium treatment but the experimentalmaterial responded markedly to light. Leaf-area ratio in thethree treatments showed extreme plasticity in increasing from2–3 x 10^–2 through 6 x 10^–2 to 8–9 x10^–2 m² g^–1 as light intensity decreased. Correspondingdecreases in unit leaf rate, however, caused over-all reductionsin relative growth rate. Specific absorption rates for potassium (A_K, dry-weight basis)were strongly reduced at the lower light intensities but alsodisplayed complex ontogenetic drifts. Values of the allometricconstant, k (the ratio of root and shoot relative growth rates),decreased from c. 0.7 at 36.1 W m^–2 through c. 0.3 at7.3 W m^–2 to a value not significantly different fromzero (P < 0.05) at 2.2 W m^–2. In material grown under the two higher light intensities a constantinverse relationship was found between the mass ratio of rootand shoot and the corresponding activity ratio. The resultsconform to this model: Mass ratio = –0.001+45.0 (1/activityratio) where activity ratio is expressed as specific absorptionrate for potassium (in µg g root^–1 h^–1)/unitshoot rate (rate of increase of whole-plant dry weight per unitshoot dry weight, in mg g shoot^–1 h^–1). The implicationsof this relationship are discussed.