Infection of Coscinodiscus spp. by the parasitoid nanoflagellate Pirsonia diadema: II. Selective infection behaviour for host species and individual host cells

The parasitoid nanoflagellate (PNF) Pirsonia diadema is hostspecific for the marine centric diatom Coscinodiscus spp. Experimentsshowed that flagellates significantly prefer C. wailesii overC.granii as host species (interspecific selectivity). This preferencewas independent of light conditions (dark, irradiance of 10and 70 µmol m^–2 s^–1) and temperature (10 and15C). Among unicellular host diatoms, the infection behaviourwas selective for individual cells: already infected C.graniicells were more attractive for further flagellate attachmentthan non-infected cells (intraspecific selectivity). Individualcells (     

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.     

Fluxes of diatoms in the Dona Paula Bay, west coast of India

Sediment traps were deployed at a station in the Dona PaulaBay to collect sedimenting particles at weekly intervals fromNovember to May during 1995–1997. Sedimented particleswere analysed for total diatom flux, chlorophyll a (Chl a) andparticulate organic carbon (POC). The highest diatom flux wasrecorded in April–May for both the years. Fluxes of diatomsvaried from0.6 x 10⁴ cells m^–2 day^–1 (November 1995)to 121.47 x 10⁴ cells m^–2 day^–1 (December 1996).In all, 19 diatom genera were identified in the sedimented material.Navicula, Nitzschia, Pleurosigma, Licmophora, Coscinodiscus,Rhizosolenia and Surirella were the most abundant genera inthe sedimented material throughout the sampling period. Meanflux of POC and diatom carbon was 251 and 0.39 mg C m^–2day^–1, respectively. The diatom carbon accounted for 0.15%of the POC flux. Mass flux of diatoms showed significant negativecorrelation with the concentration of nitrate and phosphate.This suggests that the nutrient concentration played an importantrole in influencing the sedimentation of diatoms at this coastalstation.     

Infection of Coscinodiscus spp. by the parasitoid nanoflagellate Pirsonla diadema: I. Behavioural studies on the infection process

The investigation of successive steps involved in the infectionprocess of the marine diatoms Coscinodiscus granii and Coscuwdiscuswailesii by the host-specific parasitoid nanoflagellate (PNF)Pirsonia diadenw showed that flagellates reacted chemokJinokineticallywith changes of swimming pattern to the presence of a host diatom.Chemosensory stimulation appeared to induce readiness for infection,whereas attachment and penetration of the diatom cell wall wasinduced by a mechanosensory response to morphological featureson the diatom frustules. The mean swimming speed of P.diademaflagellates decreased during their infective lifetime of 3 daysfrom an average of 78 µm s^–1 to 51 µm s^–1while the frequency of small loops in the swimming pattern increasedfrom 0.8 to 6.3 loops min^–1. At high Cgranii densities,an epidemic was delayed. It is suggested that this could becaused by overlapping gradients of extracellular material releasedby the diatoms which impaired the sensing of spatial gradientsby PNF and, therefore, the location of hosts.     

Infection of the diatom Asterionella by a chytrid. I. Effects of light on reproduction and infectivity of the parasite

The influence of hight limitation of the diatom Asterionellaformosa Hass, on the growth-determining parameters of its fungalparasiteRhizophydium planktonicum Canter emend, was measured,using laboratory cultures of both organisms. The experimentswere earned out at 6°C under a 15:9 h light-dark cycle.At saturating light conditions, the mean zoospore productionof the parasite was 23.4 zoospores sporangium^–1, and themean development time of the sporangia was 7 9 days. Light limitationof the host caused a substantial decrease of the zoospore production,while the development time was only slightly reduced. The improvedzoospore production at high light intensities was mainly theresult of incorporation of photosynthetic products generatedby the host after infection. Under limiting light conditions,Asterionella cells were less susceptible to infection withfungal zoospores. No infection at all occurred below 2 µEm^–2 s^–1, a light intensity that still supportedsome algal growth The maximum infection rate indicated thatchemotactic attraction of the parasite's zoospores by extracellularproducts of the host is involved. The infective lifetime ofthe zoospores of the parasite did not depend on light conditions,and was estimated at 8 days. The measured zoospore productionrates, both under limiting and saturating light conditions,enable the parasite to exceed the specific growth rate of thehost, and thus become epidemic, at sufficiently high host densities.     

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.     

Laboratory study of the spawning rate of the calanoid copepod Centropages typicus: effect of fluctuating food concentration

The spawning rate of laboratory-reared Centropages typicus fedHymenomonas elongala increases with food concentration, up toa value of {small tilde}2800 µg C (16 500 cells) ml^–1.An alternation of a low food (1000 cells ml^–1) and highfood concentration (16 500 cells ml^–1) is not favourableto egg release when its periodicity is 1 or 2 days, whereasit may be of advantage if it is longer (3–6 days). Inthe latter case, Centropages typicus will benefit best fromthe rich food diet if this coincides with (or just follows)the last moult.     

Seasonal dynamics in the abundance of Micromonas pusilla (Prasinophyceae) and its viruses in the Gulf of Naples (Mediterranean Sea)

To assess the role of viruses in the bloom dynamics of Micromonaspusilla in the Gulf of Naples (Mediterranean Sea), variationsof host and virus abundance were followed over one annual cycleand in late winter–spring of three consecutive years.Micromonas pusilla was recorded from autumn to spring, withpeak values up to 6.6 x 10³ cells ml^–1, but was undetectablein summer. Free M.pusilla viruses were detectable in all seasons,with concentrations from 0.02 viruses ml^–1 to 1.9 x 10³viruses ml^–1, exceeding host abundances only in one case.We found a great intraspecific variability in host susceptibilityto viruses present in natural samples, with viral titres rangingover one or two orders of magnitude for the same samples incubatedon different M.pusilla strains. Over the winter–springperiods, a highly dynamic situation was evident, with wide fluctuationsfor both host and virus abundances from one week to another.In some cases, peak host concentrations were accompanied byan increase in viral numbers, whereas in other cases the respectivefluctuations were uncoupled. Although fluctuations of M.pusillaabundance could be influenced by viral infection, there wasno evidence that viruses were able to terminate host blooms.The summer decline of M.pusilla populations did not appear tobe related to the impact of viral infection.     

The seasonal abundance of the phototrophic ciliate Mesodinium rubrum in Southampton Water, England

The abundance of the marine phototrophic planktonic ciliateMesodinium rubrum was monitored throughout an annual cycle attwo stations in the Southampton Water estuary. Seasonal changesin the concentration of nitrate, ammonia and phosphate weremonitored both at the inner estuary station (NW Netley) andouter estuary station (Calshot). Nutrient levels in the winterwere similar at both stations, and were diminished during sequentialdiatom blooms dominated initially by Sketetonema costatum andthen by Rhizosolenia dclicatula. Nitrate was reduced to a seasonalminimum in the outer estuary following these spring diatom blooms,but in the inner estuary was sustained >500 µg I^–1until the onset of the M.rubrum bloom. During the developmentof a visible red tide of M.rubrum in June/July at NW Netley,nutrient concentrations were considerably reduced. Cell numbersof M.rubrum varied between 2 and 3 cells ml^– during winterto >400 cells ml^–1 during the bloom at NW Netley, whereasat Calshot cell abundance did not increase above 25 cells ml^–1at any time of the year. At NW Netley, dense accumulations ofthe ciliate occurred over restricted depth intervals duringthe bloom and possible factors influencing the observed verticaldistribution of cells are considered. ¹Present address: Biology Department, Faculty of Science, AddisAbaba University, PO Box 1176, Addis Ababa, Ethiopia     

Growth of harmful marine algae in multispecies cultures

Mixtures of harmful and harmless algae were grown in discontinuouslydiluted batch cultures under ammonium, nitrate and phosphatelimitation, and at different irradiances (20–500 µjnolquanta m^–2 s^–1). The species used were Chrysochromulinapolylepis, Emiliania huxleyi type B, Rhodomonas sp., the dinoflagellalesFibrocapsa japonica, Gymnodinium simplex, Gyrodinium aure-olum,Heterocapsa triquetra, Heterosigma carterae, Prorocentrum micansand Alexandrium tamarense, the diatoms Chaetoceros socialis,Cymatosira belgica, Ditylum brightwellii, Laudcria borealis,Odon-telta aunla, Pseudonitzschia pungens, Streptotheca tamesis,and the cyanobacterium Synechococcus. Their growth responsein the mixed algal cultures is discussed in relation to theirabundance in different natural habitats. In comparison withthe other non-diatoms, the mixotrophic C.polylepis grew fastunder all tested nutrient and light limitations.Emiliania huxleyigrew well under nitrogen (N) limitation (with nitrate as N source)and at irradiance levels from 15 up to 500 µmol quantam^–2 s^–1. No growth of calcifying cells could bedetected under N limitation when ammonium was used as N source.Rhodomonas grew reasonably well under ammonium-N limitationand grew fast at the highest irradi-ance. The dinoflagellateswere poor competitors compared to the Prymnesiophyceae. Theenvironmental fitness of the Prymnesiophyceae appears to beclosely related to the reproductive capacity of the vegetativestage, whereas the natural distribution of dinoflagellates seemsmore closely dependent on the generative reproduction-relatedspecific life cycle characteristics of the individual species.The marine diatoms include a mixture of both types of species.Some marine diatom species clearly have the capability to outcompetenon-diatoms under different types of nutrient and light limitationswhen silicate is in excess. Other diatoms seem to be poor competitors.     

The Uptake of Nitrite by the Diatom Phaeodactylum: Interactions between Nitrite and Nitrate

Freshly harvested ammonium-grown cells of the diatom Phaeodactylumtricornutum cannot take up nitrite but they acquire the abilityto do so after a period (about 3 h) of nitrogen deprivationunder conditions that allow photosynthesis. Addition of cycloheximide(1.0 µg ml^–1) prevents development as does incubationin Na⁺ or K⁺-free medium. Nitrite uptake is dependent on thepresence of Na⁺ in the medium and is inhibited by ammonium andnitrite uptake but it is concluded that the inhibition of nitriteuptake by nitrate is not of the competitive type.     

Specific variability in Fe-enriched cultures from the equatorial Pacific

Water collected on the spring equatorial Pacific Joint GlobalOcean Flux Study (JGOFS) was placed in ultra-clean bottles inan incubator on deck under the direction of the team of J.H.Martin.Half of the bottles were enriched with 2.53 mM iron; the otherhalf served as controls. Cell counts increased, while the numberof species was reduced, in both the controls and the treatedsamples over the period of the experiment. Diatoms dominated,and after an early growth spurt in the Fe-enriched bottles,most diatom species showed greater growth in the controls atthe end of the 6 day experiment. However, as the experimentwas terminated, more cells overall were noted in the Fe-enrichedsamples, with the most abundant diatom, Cylindrotheca closterium.principally responsible for the difference. Growth rates werehigh in both the controls and the treated samples, with overallrates of 1.3–3.0 divisions day^–1 in the Fe-treatedsamples and 1.9–3.4 divisions day^–1 in the controls.The group of smaller pennate diatoms averaged 2.9 divisionsday^–1 in the Fe-treated samples and 3.3 divisions day^–1in the controls over the entire experiment.     

Composition and distribution of the pelagic and sympagic algal assemblages in the Laptev Sea during autumnal freeze-up

The phytoplankton and ice algal assemblages in the SiberianLaptev Sea during the autumnal freeze-up period of 1995 aredescribed. The spatial distribution of algal taxa (diatoms,dinoflagellates, chrysophytes, chlorophytes) in the newly formedice and waters at the surface and at 5 m depth differed considerablybetween regions. This was also true for algal biomass measuredby in situ fluorescence, chlorophyll (Chl) a and taxon-specificcarbon content. Highest in situ fluorescence and Chl a concentrations(ranging from 0.1 to 3.2 µg l^–1) occurred in surfacewaters with maxima in Buor Khaya Bay east of Lena Delta. Thealgal standing stock on the shelf consisted mainly of diatoms,dinoflagellates, chrysophytes and chlorophytes with a totalabundance (excluding unidentified flagellates <10 µm)in surface waters of 351–33 660 cells l^–1. Highestalgal abundance occurred close to the Lena Delta. Phytoplanktonbiomass (phytoplankton carbon; PPC) ranged from 0.1 to 5.3 µgC l^–1 in surface waters and from 0.3 to 2.1 µg Cl^–1 at 5 m depth, and followed the distribution patternof abundances. However, the distribution of Chl a differed considerablyfrom the distribution pattern shown by PPC. The algal assemblagein the sea ice, which could not be quantified due to high sedimentload, was dominated by diatom species, accompanied by dinoflagellates.Thus, already during the early stage of autumnal freeze-up,incorporation processes, selective enrichment and subsequentgrowth lead to differences between surface water and sea icealgal assemblages.     

Effect of temperature and food concentration on post-embryonic development, egg production and adult body size of calanoid copepod Eurytemora affinis

Post-embryonic development time, egg production rate and adultbody size of calanoid copepod Eurytemora affinis from Lake Ohnuma,Japan, were determined under six temperature-food conditions(10³,5 x 10³, 10⁴ and 5 x 10⁴ cells ml^–1 at 15°C,5 x 10⁴ cells ml^–1 at 10and20°C) in the laboratory.The measured parameters varied with both temperature and foodconcentration. Development time from hatching to adult femalewas 9.2, 11.4 and 22.8 days at 20, 15 and 10°C respectively,at the highest food concentration. The males developed to adultat one to two days earlier than the females. An effect of foodshortage on development time occurred at the lowest food concentration.This development time was 24.8 days even at 15°C, beingtwice as long as that at the highest food concentration. Prosomelength of these food-limited females was approximately 75% ofwell-fed ones, which reduced by only 10% with increasing temperaturefrom 10 to 20°C. Clutch size (C, eggs clutch^–1) ofwell-fed individuals depended on prosome length of the adultfemale (L, mm), and was expressed as an equation: C = 65.2 L³⁸³. Clutch size of individuals reared at less than 10⁴ cellsml^–1, however, mostly laid below the estimated curve,especially at the lowest food concentration being only 10% ofthat at the highest food concentration. These results suggestthat food availability is a more important factor affectingpopulation growth of E.affinis in Lake Ohnuma than variationof temperature.     

Seasonality, substrate perference and attachment sites of epizoic diatoms on cladoceran zooplankton

Cladocerans from three lakes in northwest Iowa were examinedfor attached diatoms over a period of 1 year to assess the seasonality,substrate preference and attachment site specificities of epizoicdiatoms. Over 90% of the cladocerans had attached diatoms immediatelyfollowing ice thaw, when the lakes were not yet thermally stratified,surface water temperatures were low(10–15°C)and watertransparency was at its annual high. Mean maximum density ofdiatoms on cladocerans averaged among lakes at this time was3000 cells animal^–1. The dominant epizoic diatom was Synedracyclopum Brutschy, which was found most often on large, pelagicDaphnia pulicaria and Daphnia galeata mendotae. Heavily infestedanimals had diatoms concentrated on the second antennae andanterior ventral margin of the carapace. Animals with <5epizoic diatoms were more likely to have cells attached to thepostabdomen or tail spine. In addition to living epizoically,S.cyclopum was found free living in the plankton, exhibitingtemporal fluctuations in densities similar to epizoic counterpartsHowever, the densities of the free-living cells seldom exceededepizoic densities, suggesting that S.cyclopum is adapted forgrowth on motile cladoceran substrates. ¹Present address: Savannah River Ecology Lab, Drawer E, Aiken,SC 29802, USA ²Present address: Department of Fisheries and Aquatic Sciences,7922 NW 71st St, Gainesville, FL 32606, USA     

Effect of u.v.-B irradiance on the response of 15N-nitrate uptake of Lauderia annulate and Synedra planctonica

The marine diatoms Lauderia annulata and Synedra planctonicaharvested during exponential growth were exposed to differentdoses of u.v.-B (286, 439 and 710 J m^–2 d^–1) for2 days. Uptake of ¹⁵N-nitrate was estimated before, during andafter u.v.-B radiation over 2 days. Exposure to high levelsof u.v.-B (710 J m^–2 d^–1) caused irreversible damageat the second daily irradiance. Lauderia cells were less affectedby u.v.-B stress than Synedra cells. ¹⁵N-nitrate uptake wasreduced under u.v.-B irradiance but could be reactivated within1 day following exposure to a low dose (286 J m^–2 d^–1).Higher levels of u.v.-B radiation (710 J m^–2 d^–1)led to irreversible damage. The pattern of ¹⁵N-incorporationinto several amino acids of Lauderia varied after 2 days ofu.v.-B radiation. ¹⁵N enrichment of glutamine increased markedlyafter u.v.-B stress (717 J m^–2 d^–1) whereas ^I5Nexcess of aspartic acid was significantly reduced. Results arediscussed with reference to the u.v.-B damage of the nitratetransport system.     

GROWTH OF JUVENILE BITHYNIA TENTACULATA (PROSOBRANCHIA, BITHYNIIDAE) UNDER DIFFERENT FOOD REGIMES: A LONG-TERM LABORATORY STUDY

The growth of juvenile Bithynia tentaculata (Proso-branchia,Bithyniidae) was measured under controlled laboratory conditionsover 18 months. The animals were fed with different concentrationsof suspended food (Chlamydomonas reinhardii at 2, 000 cells.ml^–1and at 10, 000 cells.ml^–1), solid food (lettuce) and combinationsof both (lettuce with 2, 000 cells.ml^–1 and with 10, 000cells.ml^–1 Chlamydomonas). The growth of all animals wasmeasured weekly, and after 1 years final shell sizes, shelldry weights and ash-free dry weights, as well as soft body dryweights and ash-free dry weights were determined. Survival rateof the animals increased from 20% when fed with 2, 000 algalcells.ml^–1 to 75% with lettuce and 10, 000 cells.ml^–1as food. Final shell sizes and shell weights were not influencedby food, but soft body dry weights were significantly reducedwhen animals were fed on suspended food only. Reproduction (i.e.egg laying) was observed at the end of the second summer whenlettuce (with or without algal addition) was given as food.The ecological consequences of these results are discussed. (Received 7 April 1993; accepted 9 August 1994)     

Hyperscums of the cyanobacterium Microcystis aeruginosa in a hypertrophic lake (Hartbeespoort Dam, South Africa)

It is proposed that surface scums of densely packed planktoniccyanobacteria (blue-green algae) which exist for weeks to months,measure several decimeters in thickness and are covered by acrust of photo-oxidized cells, be called hyperscums. Hyperscumsof Microcystis aeruginosa formed during prolonged periods ofcalm weather in wind-protected sites in a hypertrophic lakesubject to low wind speeds (Hart beespoort Dam, South Africa).A hyperscum that extended over 1–2 hectares and persistedfor 103 days during winter 1983 was studied. Chlorophyll a concentrationsranged from 100 to 300 mg l^–2 Microcystis cell concentrationsreached 1.76x10⁹ cells ml^–1 or 116 cm³l^–1. The hyperscumenvironment was anoxic, aphotic, with a fluctuating temperatureregime and low pH values. The densely packed Microcystis cellssurvived these conditions for more than 2 months. This was shownby comparing the potential photosynthetic capacity of Microcystisfrom the hyperscum with that of Microcystis from the main basinof the lake. However, after 3 months the hyperscum algae losttheir photosynthetic capacity and decomposition processes prevailed.The hyperscum gradually shrank in size until a storm causedits complete disintegration.     

Buoyancy properties of the giant diatom Ethmodiscus

Net-collected Ethmodiscus spp. were examined and found to becapable of positive buoyancy with ascent rates averaging 1.4m h^–1 on individually measured cells, and 0.3 m h^–1using a homogeneous settling technique. These data, togetherwith published reports of positive buoyancy in Rhizosolenia,indicate that upward movement is a widespread phenomenon inlarge oceanic diatoms. These observations also suggest thatpublished size-rate relationships for marine diatoms requirerevision to include the effects of positive buoyancy.     

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.