Laboratory and field observations on the scuticociliate Histiobalantium from the pelagic zone of Lake Constance, FRG

Histiobalantium sp. was found regularly in the pelagic zoneof Lake Constance, FRG, over five annual cycles. Maxima of upto 6400 cells l^–1 were recorded in late summer, with similarnumbers in the 0–8 and 8–20 m depth intervals. Onan annual average, the population accounted for 10–17%of the total biomass of planktonic ciliates. In the laboratory,Histiobalantium grew well on a diet of the cryptophyte Rhodomonassp. Maximum growth rates obtained in batch cultures were 0.21and 0.33 day^–11 at 9 and 18°C, respectively. In situexperiments using diffusion chambers yielded positive growthrates in autumn and winter. The highest values recorded at theambient temperatures 5, 14 and 17°C were 0.17, 0.32 and0.40 day^–1, respectively. Comparing these results withthe different seasonal distributions and higher measured growthrates of other ciliates from Lake Constance, we conclude thatHistiobalantium is a superior competitor at relatively low algalfood concentrations. ²Present address: Fisheries & Oceans Canada, 4160 MarineDrive, West Vancouver, BC, V7V 1N6, Canada     

Predation impact of Cyclops vicinus on the rotifer community in Lake Constance in spring

The predation impact of Cyclops vicinus on rotifers was studiedunder near-natural conditions in small enclosures to evaluatewhether copepod predation is responsible for the decline ofrotifers in Lake Constance in spring. Cyclops vicinus fed selectivelyon Synchaeta spp.; Keratella and Polyarthra spp. were not selectedfor. Predation rates increased with prey density up to a maximumof 37 Synchueta day^–1 at a density of 1.6 x 10⁶ Synchaetam^–2, i.e. at -1200 Synchaeta l^–1. Calculation ofcropping rates suggests that Cyclops alone can control the abundanceof Synchaeta in spring, i.e. that mainly Cyclops is responsiblefor the decline of Synchaeta species in Lake Constance in May.     

Comparative growth rates and yields of ciliates and heterotrophic dinoflagellates

Growth rates, ingestion rates and grazer yields (grazer volumeproduced/prey volume consumed) were measured for six protozoanspecies (ciliates: Favella sp., Strombidinopsis acuminatum,Uronema sp.; heterotrophic dinoflagellates: Amphidinium sp.,Gymnodinium sp., Noctiluca scintillans) in laboratory batchculture experiments. Comparative growth data indicate that theprymnesiophyte Isochrysis galbana, the prasinophyte Mantoniellasquamata, two cryptophyte species and several autotrophic dinoflagellatespecies were suitable foods for these grazers. When grown onoptimized diets at 13C, maximum ciliate growth rates (range0.77–1.01 day^–1 uniformly exceeded maximum heterotrophicdioflagellate growth rates (range 0.41–0.48 day^–1).A compilation of published data demonstrates that this growthrate difference persists across a range of ciliate and dinoflagellatetaxa and cell sizes. Comparison of volume-specific ingestionrates and yields for the six species studied here showed thatthere was no single explanation for this growth rate disparity.Heterotrophic dinoflagellates exhibited both low ingestion ratesand, in one case, low yields; ciliates were able to achievehigher growth rates via either higher ingestion rates or higheryields, depending on ciliate species. Volume yield increasedover time throughout the exponential growth phase in nearlyall experiments, suggesting variation in response to changingfood concentrations or long-term acclimation to culture conditions.Higher maximum ciliate growth rates mean that these grazershave the potential to exercise tighter control over incipientblooms of their prey than do heterotrophic dinoflagellates.     

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.     

Development, growth and egg production of Centropages abdominalis in the eastern subarctic Pacific

Centropages abdominalis is a neritic, omnivorous, temporallyabundant copepod present throughout the subarctic Pacific andits marginal seas. The two main objectives of this study wereto determine how temperature influences the development of C.abdominalis and whether growth rates of in situ populationsmay be limited by available food. At 6.9°C, median developmenttime from eggs laid to 50% adults was 42 days and the averageweight-specific somatic growth rate was 0.17 day^–1. At4.6°C, median development time to adult was 59 days (projected)and growth rate averaged 0.08 day^–1, suggesting that 4.6°Cmay be approaching the lower temperature for development andgrowth in this species. The functional relationship betweendevelopment time and temperature was established over the temperaturerange in which this species occurs. The in situ adult growthrates between 10 and 13°C averaged 0.14 day^–1 andwere generally lower than the laboratory-reared juvenile growthrates, which may indicate that adult C. abdominalis are foodlimited in the field during summer and autumn.     

Growth and development of Neocalanus flemingeri/plumchrus in the northern Gulf of Alaska: validation of the artificial-cohort method in cold waters

In situ growth and development of Neocalanus flemingeri/plumchrusstage C1–C4 copepodites were estimated by both the artificial-cohortand the single-stage incubation methods in March, April andMay of 2001–2005 at 5–6°C. Results from thesetwo methods were comparable and consistent. In the field, C1–C4stage durations ranged from 7 to >100 days, dependent ontemperature and chlorophyll a (Chl a) concentration. Averagestage durations were 12.4–14.1 days, yielding an averageof 56 days to reach C5, but under optimal conditions stage durationswere closer to 10 days, shortening the time to reach C5 (fromC1) to 46 days. Generally, growth rates decreased with increasingstage, ranging from 0.28 day^–1 to close to zero but weretypically between 0.20 and 0.05 day^–1, averaging 0.110± 0.006 day^–1 (mean ± SE) for single-stageand 0.107 ± 0.005 day^–1 (mean ± SE) forartificial-cohort methods. Growth was well described by equationsof Michaelis–Menten form, with maximum growth rates (G_max)of 0.17–0.18 day^–1 and half saturation Chl a concentrations(K_chl) of 0.45–0.46 mg m^–3 for combined C1–3,while G_max dropped to 0.08–0.09 day^–1 but K_chl remainedat 0.38–0.93 mg m^–3 for C4. In this study, in situgrowth of N. flemingeri/plumchrus was frequently food limitedto some degree, particularly during March. A comparison withglobal models of copepod growth rates suggests that these modelsstill require considerable refinement. We suggest that the artificial-cohortmethod is the most practical approach to generating the multispeciesdata required to address these deficiencies.     

Pseudobalanion planctonicum (Ciliophora, Prostomatida): ecological significance of an algivorous nanociliate in a deep meso-eutrophic lake

Pseudobalanion plancioracum was the most abundant ciliate speciesin the pelagic zone of Lake Constance, FRG, over a 3 year period.Annual averages accounted for 30, 35 and 43% of total ciliatenumbers in the uppermost 20 m of the water column in 1987, 1988and 1989 respectively. Highest cell numbers were observed inearly spring, simultaneously with the first phytoplankton maximum.The small ciliate (mean length 15 µm, mean cell volume1300 µm³) is a raptorial feeder and predominantly consumesphytoplankton. In laboratory cultures, P planctonicum grew wellon a diet of Rhodomonas sp. (Cryptophyceae). Maximum growthrates increased from 0.46 day^–1 at 5 5°C to 1.52 day^–1at 18.5° C, while temperatures above 21°C were lethal.Depending on food concentration, 0.2–4.4 Rhodomonas cellswere ingested per ciliate and hour.     

Specific growth rates of heterotrophic plankton organisms in a eutrophic lake during a spring bloom

The in situ growth of the dominating pelagic organisms at severaltrophic levels was investigated during a spring bloom characterizedby well-mixed cold water. The study includes primary productionand the carbon flow through the nano-, micro- and mesozooplanktonpopulations based on population dynamics and specific growthrates. The phytoplankton biomass and production were totallydominated by small algae <20 µm. of which {small tilde}5%were <3µm. potentially a food source for the nano-and microzooplankton. The mean carbon-specific primary productionwas 0.15 day^–1 and was regulated solely by light. Themean volume-based specific growth rate of bacterioplankton wasmodest. 0.1 day^–1. and probably controlled by the lowtemperature. The volume-based specific growth rates of heterotrophicnanoflagellates. ciliates. rotifers and copepods were 0.35.0.13. 0.16 and 0.03 day^–1, respectively. The observedgrowth of the heterotrophic plankton was generally not foodlimited, but was controlled by temperature. The stable temperatureduring the experiment therefore allows a cross-taxonomic comparisonof specific growth rates. The b exponent in the allometric relationship(G = aV^th) between volume-specific growth rate (G) and individualbody size (V) was –0.15 ± 0.03 for all filtratingzooplankton. indicating an in situ scaling not far from thephysiological principles onginally demonstrated for laboratorypopulations.     

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

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

On temperature-independent growth of phytoplankton

Growth rates of two freshwater diatoms and three chlorophyteswere compared under 3 h days at 10 µmol m^–2 s^–1.Specific growth rates ranged between µ=0.03 day^–1and µ=0.055 day^–1 for the different species andwere in every species independent of temperature between 8 and16°C.     

Growth and decline of a diatom spring bloom phytoplankton species composition, formation of marine snow and the role of heterotrophic dinoflagellates

During the spring of 1994, we determined the factors responsiblefor the decline of the seasonal diatom bloom in the Gullmarfjord, on the west coast of Sweden. Four species constituted>75% of the biomass—Detonula confervacea, Chaetocerosdiadema, Skeletonema costatum and Thalassiosira nordenskioeldii—reachingconcentrations of 4900, 350, 8200 and 270 cells ml^–1,respectively. Growth of phytoplankton was exponential (growthrate = 0.12 day^–1) from 3 to 21 March, after which a galewith winds >15 m s^–1 caused massive aggregation. Amaximum of 130 p.p.m. (v/v) of marine snow aggregates was observedby in situ video at the peak of the bloom. Critical concentrations(Jackson, Deep-Sea Res., 37, 1197–1211, 1990) were similarto observed showing that coagulation theory could explain thesudden decline of the bloom. The heterotrophic dinoflagellateGyrodinium cf. spirale increased exponentially after the peakof the bloom with maximum (temperature-adjusted) growth rates.After the rapid aggregation and sedimentation of the bloom,they were able to control any further growth of diatoms. Nitrateand silicate were never depleted, but phosphate may have beenlimiting by the end of the study period. We conclude that massaggregation during a gale marked the end of the bloom, and thatintense grazing by heterotrophic dinoflagellates prevented anysubsequent increase of diatoms.     

In situ determinations of bacterial selectivity and filtration rates by five cladoceran zooplankters in a hypertrophic subtropical reservoir

Cladoceran in situ feeding rates on natural bacteria labelledwith [methyl-³H] were studied in parallel with feeding ratedeterminations on ¹⁴C-labelled Chlorella in a hypertrophic subtropicalreservoir (Lake Hartbeespoort) through spring and summer (1986/87).Community filtration rates (CFR5) on bacteria and algae weresimilar, but selection for Chlorella (relative to natural bacteria)increased from midsummer in association with declining bacterialdensity and increasing dominance of ‘inedible’ componentsof the natural phytoplankton. Species-specific filtration rates(SSFRs) were determined for Daphnia pulexllongispina, Ceriodaphniareticulata, Diaphanosoma excisum, Bosmina longirostris and Moinamicrura during their respective seasonal occurrence in the studyperiod. SSFRs on algae and bacteria increased with body length(L, mm) in all species apart from Bosmina. Species-specificdifferences in absolute feeding rate (FR, ml animal^–1day^–1), the slope of the FR-L relationship and bacteriaselectivity were evident. The feeding rate of all cladoceranson bacteria is described by the power equation FR 5.231L^1.42FR values on bacteria relative to FR values on algae averaged     

Dynamics of microplankton communities at the ice-edge zone of the Lazarev Sea during a summer drogue study

Microzooplankton grazing and community structure were investigatedin the austral summer of 1995 during a Southern Ocean Drogueand Ocean Flux Study (SODOFS) at the ice-edge zone of the LazarevSea. Grazing was estimated at the surface chlorophyll maximum(5–10 m) by employing the sequential dilution technique.Chlorophyll a concentrations were dominated by chainformingmicrophytoplankton (>20 µm) of the genera Chaetocerosand Nitzschia. Microzooplankton were numerically dominated byaloricate ciliates and dinoflagellates (Protoperidinium sp.,Amphisoleta sp. and Gymnodinium sp.). Instantaneous growth ratesof nanophytoplankton (<20 µm) varied between 0.019and 0.080 day^–1, equivalent to between 0.03 and 0.12 chlorophylldoublings day^–1. Instantaneous grazing rates of microzooplanktonon nanophytoplankton varied from 0.012 to 0.052 day^–1.This corresponds to a nanophytoplankton daily loss of between1.3 and 7.0% (mean = 3.76%) of the initial standing stock, andbetween 45 and 97% (mean = 70.37%) of the daily potential production.Growth rates of microphytoplankton (>20 µm) were lower,varying between 0.011 and 0.070 day^–1, equivalent to 0.015–0.097chlorophyll doublings day^–1. At only three of the 10 stationsdid grazing by microzooplankton result in a decrease in microphytoplanktonconcentration. At these stations instantaneous grazing ratesof microzooplankton on microphytoplankton ranged between 0.009and 0.015 day^–1, equivalent to a daily loss of <1.56%(mean = 1.11%) of initial standing stock and <40% (mean =28.55%) of the potential production. Time series grazing experimentsconducted at 6 h intervals did not show any diel patterns ofgrazing by microzooplankton. Our data show that microzooplanktongrazing at the ice edge were not sufficient to prevent chlorophylla accumulation in regions dominated by rnicrophytoplankton.Here, the major biological routes for the uptake of carbon thereforeappear to be grazing by metazoans or the sedimentation of phytoplanktoncells. Under these conditions, the biological pump will be relativelyefficient in the drawdown of atmospheric CO₂.     

The relative importance of food and temperature to copepod egg production and somatic growth in the southern Benguela upwelling system

The fecundity and somatic growth rates of Calanus agulhensisand Calanoides carinatus, the dominant large calanoid copepodsin the southern Benguela upwelling system, as well as the fecundityof several other common copepods, were measured between Septemberand March of 1993/94 and 1994/95. Mean egg production of mostcopepods was low at >30 eggs female^-1 day^-1 {Calanoides carinatus23.7, Calanus agulhensis 19.0, Neocalanus tonsus 16.1 and Rhincalanusnasutus 26.1), whereas the mean fecundity of Centropages brachiatuswas significantly greater (83.6 eggs female^–1 day^-1).This study also presents the first comprehensive field estimatesof the fecundity of Nanno-calanus minor (mean: 26.1 eggs female^–1day^–1, range: 0.0–96.2 eggs female^–1 day^–1)and of somatic growth of N6 and all copepodite stages of Calanoidescarinatus (decreasing from 0.58 day^–1 for N6 to 0.04 day^–1for C5). Somatic growth rates of Calanus agulhensis also declinedwith age: from 0.57 day1 for N6 to 0.09 day1 for C5. Data ongrowth rates were used to assess the relative importance offood [as measured by total chlorophyll (Chi) a concentration],phytoplankton cell size (proportion of cells >10 µm)and temperature to the growth of copepods. Multiple regressionresults suggested that fecundity and somatic growth rates werepositively related to both Chi a concentration and phytoplanktoncell size, but not to temperature. Although it was not possibleto separate the effects of Chi a concentration and phytoplanktoncell size, data from previous laboratory experiments suggestthat copepod growth is not limited by small cells per se, butby the low Chi a concentrations that are associated with theseparticles in the field. Despite growth not being directly relatedto temperature, a dome-shaped relationship was evident in somespecies, with slower growth rates at cool (<13°C) andwarm (>18°C) temperatures. The shape of this relationshipmirrors that of Chi a versus temperature, where poor Chi a concentrationsare associated with cool and warm temperatures. It is concludedthat the effect of food limitation on growth of copepods outweighsthat of temperature in the southern Benguela region. Sourcesof variability in relationships between growth and Chi a concentrationare discussed.     

Zooplankton growth rates: extraordinary production by the larvacean Oikopleura dioica in tropical waters

The growth of Oikopleura dioica was determined from microcosmsincubated in situ at 29°C in Kingston Harbour, Jamaica.Minimum generation times approached 1 day, with Oikopleura dailyspecific growth rates ranging from 3 to 23 and averaging 10.7-foldincreases in biomass over 24 h. This was an order of magnitudegreater than the copepod Paracalanus crassirostris, whose growthrates ranged from 0.3 to 1.2 and averaged 0.67 day^–1 fromthe same experiments A reassessment of previous data indicatesthat larvacean production approaches, and may exceed, the 693kJ m^–2 annual production of the copepods. Growth rateand recruitment of Oikopleura decreased as the biomass of thezooplankton community increased; both resource and interferencecompetition are probably occurring. The extraordinary productionpotential of the Larvacea emphasizes their understated importancein marine planktonic systems.     

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     

Feeding and metabolism of the siphonophore Sphaeronectes gracilis

The in situ predation rate of the siphonophore Sphaeronectesgracilis was estimated from gut content analysis of hand-collectedsiphonophores and from laboratory data on digestion rates ofprey organisms. At daytime prey densities of 0.25 copepods 1^–1,S. gracilis was estimated to consume 8.1 – 15.4 prey day^–1siphonophore^–1. From data on abundances of siphonophoresand copepods, S. gracilis was estimated to consume 2–4%of the copepods daily. In laboratory experiments, ingestionrates averaged 13.8 prey day^–1 siphonophore^–1 atprey densities of 5 copepods 1^–1 and 36.9 at 20 copeods1^–1. This was equivalent to a specific ingestion rate(for both carbon and nitrogen) of –17% day^–1 and45% day^–1, respectively, while specific ingestion in situwas only 2% day^–1. Ammonium excretion averaged 0.095 µg-atsiphonophore^–1 day^–1 at 5 prey 1^–1, and 0.162at 20 prey 1^–1. The specific respiration (carbon) andspecific excretion (nitrogen as ammonium) were calculated tobe 3% day^–1 at the lower experimental food level, and5% day^–1 at the higher food level. ¹Contribution from the Catalina Marine Science Center No. 66. ²Present address: Dept. of Biology, University of Victoria,Victoria, B.C., Canada V8W 2Y2.     

In situ filtration responses of Daphnia galeata to changes in food quality

In the stoichiometric study of phyto–zooplankton interactions,a controversy exists about how Daphnia species regulate theirfeeding rate when submitted to low-quality food (i.e. high dietarycarbon:phosphorus [C:P] ratio). In this study, we gathered dataover 3 years on in situ clearance rates of a Daphnia galeatapopulation, by conducting grazing experiments from April 1998to October 2000 in the Esch-sur-Sûre reservoir (Grand-Duchyof Luxembourg). Observed clearance rates (2.5–13.5 mLindividual^–1 day^–1, mean 7.0 mL individual^–1day^–1) were correlated with population and environmentalvariables. Mean body size of Daphnia individuals was the bestpredictor of clearance rate (r² = 0.639), followed by watertemperature (r² = 0.262) and P concentration in the seston (r²     

Daily irradiance governs growth rate and colony formation of Phaeocystis (Prymnesiophyceae)

Phaeocystis was cultured at a range of ecologically significantdaily irradiances under nutrient-replete conditions. Below athreshold of 100 W h m^–1 day^–1, the cells were smalland flagellated, and remained solitary. Above this threshold,the cells were larger and able to form colonies. Growth rateand colony formation were maximum at sea surface irradiances(>700 W h m^–2 day^–2). Presumably, colonial growthis a strategy to maintain optimum growth rates in the watercolumn. Sinking, nutrient-stressed colonies reach low irradiancesand colonial cells can transform into small solitary flagellatedcells. These observations are important in understanding theecology and life cycle of Phaeocystis.     

Response of ciliates and Cryptomonas to the spring cohort of a cyclopoid copepod in a shallow hypereutrophic lake

The impact of a cyclopoid copepod population on the protozoacommunity (two ciliate categories and Cryptomonas) was assessedweekly during the spring cohort of Cyclops vicinus (one monthduration) in hypereutrophic Lake Søbygård by insitu gradient experiments with manipulation of ambient zooplanktonabundance. As C.vicinus always made up >92% of the zooplanktonbiomass, the response of protozoa is assumed to be a resultof predation by the copepod. Significant effects of copepodbiomass on protozoa net population growth rates were obtainedin the four experiments. Copepod clearance rates were significantlyhigher on oligotrichs than on prostomatids and Cryptomonas butdeclined for all three protozoa categories during the firstthree weeks of the copepod cohort, probably because of the changein developmental instar composition of the copepod population.Grazing impact on protozoa at ambient copepod abundance wasconsiderable (range, 0.05–0.87 day^–1) and could,together with the estimated reproductive potential of protozoans(range, –0.20–0.87 day^–1), account for thedecline in abundance and biomass of protozoa during the cohortdevelopment. Carbon flow from the protozoa to C.vicinus (range,2.8–23.5 µg C l^–1 day^–1) documents thepresence of a trophic link between protozoa and the spring cohortof C.vicinus in Lake Søbygård.