Dynamics of Copepod Faecal Pellets in Relation to a Phaeocystis Dominated Phytoplankton Bloom: Characteristics, Production and Flux

Copepod faecal pellet characteristics and production were measuredin spring 1995, 1996 and 1997 in the North Sea Southern Bightin order to define changes due to the development of the phytoplanktonbloom. Changes were related to the succession from diatomsto a Phaeocystis sp. bloom. Mean monthly pellet productiondecreased during the Phaeocystis bloom development to 0.27 pelletsind. ^–1 h ^–1, approximately 80% lower than beforeand after the bloom. Although phytoplanktonic pigments showedsignificant differences between inshore and offshore stations,there was no such significant difference for faecal pellet production.Faecal pellet sinking-rate decreased from 100 to 70 m day ^–1during the transition from a diatom- to a Phaeocystis-dominatedbloom. This was due to a decrease in pellet density and/or adecrease of the pellet volume. These results supported the ideaof lower feeding of copepods under Phaeocystis bloom conditions.As mean seasonal density of copepod faecal pellets was higher(1.37 g cm ^–3) than in other seas, accidental ingestionof sedimentary material as the cause of this high density isdiscussed.     

Changes in the elemental composition of Asterionella formosa during the diatom spring bloom

Changes in the elemental composition of the diatom Asterionellaformosa within mixed phytoplankton samples were determined overthe spring bloom period using energy dispersive X-ray microanalysis.Using a 10 kV electron microprobe, X-ray information from thetop 1–2 µm of the cell revealed overall mean concentrationsof: Si (4636 mmol kg^–1 dry wt), P (82), S (54), Cl (71)K (94) and Ca (90). Concentrations of all elements showed widevariation within each date sample, with unimodal frequency distributionsapproximating to a Normal distribution. Correlation of the entiredata set demonstrated clear statistical associations betweenelements, including Si, P and K. Uptake of Si during the courseof the diatom bloom led to a major fall in lake water concentration(1.0 to 0.07 mg l^–1), which correlated with a decreasein the mean cell Si concentration (6735 to 3107 mmol kg^–1dry wt) during the final phase of the bloom. Mean cell concentrationsof P and K also showed marked decreases with time, in closeparallel with cell Si. These changes in P and K were attributedto the high level of internal correlation with Si and not toany significant decrease in P and K availability in the environment.     

MOLECULAR EVIDENCE IDENTIFIES BLOOM-FORMING PHAEOCYSTIS (PRYMNESIOPHYTA) FROM COASTAL WATERS OF SOUTHEAST CHINA AS PHAEOCYSTIS GLOBOSA

Chen, Y. Q., Zhou, H. & Qu, L. H. Key Laboratory of Gene Engineering of Education Ministry, Biotechnology Research Center, Zhongshan University, Guangzhou 510275, P. R. China Sequence data from the 18S small subunit ribosomal RNA gene and rDNA ITS regions have been used to identify the species of a Phaeocystis (Prymnesiophyta) that caused harmful algae blooms in the coastal waters of southeast China. This Phaeocystis has morphological and physiological features that differ from those previously described for either P. globosa Scherf- fel or P. pouchetii (Hariot)Lagerheim. However, the sequence comparison of the Phaeocystis 18S rDNA and rDNA ITS clearly showed that it was remarkably similar to several isolates of P. globosa. Thus, the species isolated from the southeast coast of China is identified as P. globosa rather than P. cf. pouchetii or another species. Our results also demonstrate that phenotypes of different members of the genus Phaeocystis are variable, apparently changing in response to environmental conditions. It is concluded that, on the basis of this phylogenetic analysis, the bloom forming southeast China coast species of Phaeocystis most likely originated from an endemic warm-water, rather than a foreign source.     

Dinophysis blooms in Greek coastal waters (Thermaikos Gulf, NW Aegean Sea)

The first documented Dinophysis bloom from Greek coastal watersassociated with a diarrhetic shellfish toxins outbreak was recordedin January 2000 during the HAB monitoring program in ThermaikosGulf. A species with morphological features similar to D. acuminataClaparède and Lachmann dominated this bloom. MaximumD. cf. acuminata abundance (85.4 x 10³ cells L^–1) appearedin February 2000. In 2001, high numbers of D. cf. acuminata(5 x 10³ cells L^–1) were recorded in April under conditionsof weak thermal stratification, while in 2002 maximum numbers(37 x 10³ cells L^–1) of the same species were found inFebruary under low temperature levels (11.5–12.5°C).All Dinophysis blooms persisted for no more than 4 months. Inaddition to cells resembling typical D. cf. acuminata, cellsof similar shape but smaller in size were observed at differentstages of the blooms. Other observations include couplets oflarge and small-sized cells, and also a larger robust form witha dense granular cytoplasm. The D. cf. acuminata populationalways presented a stratified vertical distribution with verticalpeaks positioned in or just above the pycnocline. Among thephysico-chemical parameters, water temperature appears to bethe most important factor influencing the distribution of Dinophysisabundance.     

Microbial dynamics during the decline of a spring diatom bloom in the Northeast Atlantic

The microbial dynamics during a spring diatom bloom declinewas monitored in the Northeast Atlantic during a 5-day Lagrangianstudy (8–12 April 2002). Phytoplankton abundance, compositionand health status were related to viral and bacterial abundance,zooplankton abundance and grazing rates, as well as bacterialproduction. Phytoplankton reached maximum concentration on Day3 (Chl a >5 µg L^–1) and declined on Day 5 (Chla 2 µg L^–1) and was dominated (70% of Chl a) bydiatoms. Bacterial production increased substantially to >20µg C L^–1 day^–1 on Day 3 and concomitantlylarge viruses decreased in number by half to <10 x 10³ mL^–1.This was followed by a 5-fold increase in large viruses on Day5, indicating infection and subsequent lysis on Days 3 and 5,respectively. Micro- and mesozooplankton grazing were not theprincipal cause for the decline of the bloom and pheophorbide-ashowing little variation in concentration from Days 1–4(100 ng L^–1) although doubled on Day 5. The poor physiologicalstatus of the diatoms, indicated by the high chlorophyllide-aconcentrations (50–480 ng L^–1), likely promoteda series of closely interrelated events involving bacteria andviruses leading to the demise of the diatom bloom.     

Relationship between vegetative cells and cyst production during Alexandrium minutum bloom in Arenys de Mar harbour (NW Mediterranean)

A recurrent Alexandrium minutum bloom in the Arenys de Mar harbour(Catalan coast, North Western Mediterranean) was monitored inorder to establish the relationship between vegetative cellsand cyst production. The bloom lasted from January 21 to February24, 2002 and reached cell concentrations of up to 47 x 10⁶ cellL^–1. Two aspects related to the resting cysts depositionwere studied: (i) production of resting cysts during the bloomperiod (by means of sediment traps) and (ii) distribution ofresting cysts in the sediment after the bloom (May 2002). Cystformation in Arenys clearly started in a period with high vegetativecell densities in the water column. Once production was initiatedencystment fluxes remained constant for two weeks, and coveringthe periods of maintenance and decline of the bloom. High cystfluxes (up to 6000 cysts cm^–2 day^–1) were quantifiedas a result of the high vegetative cell concentration. Moreover,encystment occurring in less than 1% of the total populationindicates that most of the cells are not involved in restingcysts formation. A comparison of the resting cyst flux valuesobtained from the sediment traps and the resting cyst concentrationsin surface sediment (628–3270 cysts cm^–3) threemonths later, revealed that the number of cysts in the sedimentdecreased during that time. The studies of excystment showeda high germination percentage (91%) and germling viability (100%).These data, together with the resting cyst distribution in thesediment, are important in assessing the role of resting cystsin the bloom dynamics of A. minutum in confined waters.     

Pigment signatures reveal temporal and regional differences in taxonomic phytoplankton composition off the west coast of Ireland

The composition of phytoplankton assemblages in April, 1998in Galway Bay and during a summer phytoplankton bloom occurringsouthwest of Ireland in August, 1998, was characterized by pigmentsmeasured by high-performance liquid chromatography. Pigmentdata reflecting phytoplankton assemblages dominated by diatomsin Galway Bay and dinoflagellates in the southwest of Irelandwere compared to phytoplankton cell counts. Significant relationshipswere found between the fucoxanthin concentrations and the diatomcell numbers (P < 0.0002 r² 0.63) during April, and betweenfucoxanthin (P < 0.0001 r² 0.79), 19'hexanoyloxyfucoxanthin(P < 0.0001 r² 0.77) concentrations and Gyrodinium aureolumcell numbers during the summer bloom.     

Planktonic nitrogen transformations during a declining cyanobacteria bloom in the Baltic Sea

The uptake of ¹⁵N-labelled nitrogen nutrients (ammonium, urea,nitrate) was studied during the decline of a bloom of nitrogen-fixingcyanobacteria in the Baltic Sea. This was done by sampling anorth-south transect of stations, representing different stagesof the bloom. Comparison with nitrogen fixation data showedthat this process was of minor importance, and that the nitrogenuptake was dominated by regenerated nitrogen, mainly ammonium.From time series incubations for studying nutrient uptake, itappears that the regeneration of ammonium was substantial, butthat the production of urea or nitrate was slow. The integrateddaily uptake was calculated for the 0–15 m interval atfour stations and values ranged between 6 and 21 mmol N m^–2day^–1, of which the regenerated nutrients, ammonium andurea, constituted 71–93%. Nitrate was of minor importanceand the highest nitrate uptake rates were found close to thethermocline (at 15 m) and in the southern part of the Baltic.Comparison with carbon fixation data reported from simultaneousmeasurements at two stations gave C/N uptake ratios of 4.9 and2.1 for integrated daily uptake. Contrary to earlier findings,the concentration of DON increased with increasing salinity(from 15 to 17 µmol l^–1). This was correlated withthe declination of the bloom and is suggested to be a resultof a gradual release of less easily utilized DON from the degradationof cyanobacteria. The C/N ratio of DOM was high, 21–23.     

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.     

Cyst formation: an important mechanism for the termination of Scrippsiella trochoidea (Dinophyceae) bloom

A sediment trap study was conducted at Daya Bay, South ChinaSea, to investigate the relationships between encystment andpopulation dynamics of Scrippsiella trochoidea from December1999 to January 2001. A dense bloom of S. trochoidea occurredduring the study period from August to September 2000, withthe maximum cell number of 3.18 x 10⁴ cells mL^–1.Two morphotypes of cysts, one with a thick calcareous wall (calcifiedcyst) and another without the obvious calcareous cover (non-calcifiedcyst), were observed during this investigation. The morphologicaland excystment characteristics of these two cyst types werestudied as well. Mass encystments of S. trochoidea, with themaximum of 3.05 x 10⁵ cysts m^–2 d^–1for calcified cyst, and 1.54 x 10⁷ cysts m^–2 d^–1for non-calcified cyst, coincided with the maximum abundanceof the vegetative cells. Encystment caused the transfer of atotal of 2.24–4.49 x 10⁸ cells m^–2 vegetativecells from the water column to the sea bottom during the bloomand resulted in a considerable loss of the bloom population.High assemblages of cysts of S. trochoidea were detected inthe surface sediments as well. This rich ‘seed bed’in the surface sediments caused by the high efficiency of encystmentafter blooms acting as a benthic reservoir for future vegetativepopulation, together with the short dormant period (15–26days) and high germination rate (50–90%), may explainthe repeated occurrence of S. trochoidea blooms in Daya Bay.     

Grazing and colony size development in Phaeocystis globosa (Prymnesiophyceae): the role of a chemical signal

The bloom-forming prymnesiophyte Phaeocystis globosa forms hollow,spherical, mucilaginous colonies that vary from micrometresto millimetres in size. A recent paper gave the first empiricalevidence that colony size increase in P. globosa is a defensiveresponse against grazers, and knowing the signalling mechanism(s)behind this response will thus be a key to understanding thetrophodynamics in systems dominated by this species. I conductedexperiments with specially designed diffusion incubators, eachof which consists of a non-grazing chamber (with P. globosaonly) and a grazing chamber (grazers + phytoplankton) connectedby 2 µm polycarbonate membrane filters. The results showedthat physical contact with grazers was not required to initiatethe defensive response; instead, P. globosa colony size increasewas found to be stimulated by dissolved chemicals generatedby ambient grazing activities. This signal was non-species specific,such that various combinations of three species of grazers andfour species of phytoplankton in the grazing chambers all resultedin significant, but different extents of colony enlargementin P. globosa in the non-grazing chambers (30–300% largerthan the ‘grazer-free’ control). High concentrationsof ambient solitary P. globosa cells and other phytoplanktonseemed to suppress colony enlargement in P. globosa, and grazerswould help reduce this inhibition by removing the ambient solitaryP. globosa cells and other phytoplankton. These non-species-specificmechanisms would allow P. globosa to regulate colony size developmentand defend itself in diverse planktonic systems, which may helpto explain the global success of this species.     

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     

The seasonal population changes and carbon budget of the calanoid copepod Boeckella minuta Sars in a newly formed sub-tropical reservoir

The population carbon budget and seasonality of Boeckella minutain a newly formed subtropical reservoir were examined 3 yearsafter the reservoir filled. Average daily biomass was 26.4 mgC m^–3 and the annual population carbon budget was: consumption2470, egestion 1482, assimilation 988, production 493 and respiration495, mg C m^–3 year^–1, and the average P/B and P/Aratios were 0.08 and 0.5 respectively. Clutch size and reproductiveeffort (egg production/assimilation) were low, and the proportionof males decreased throughout the population cycle. The seasonalabundance pattern changed from perennial (pre-filling years)to a 7 month cycle. It is suggested that eutrophication andthe spring bloom of cyanobacteria may have accentuated a seasonaldecrease in reproductive effort and survival, leading to anabsence of planktonic stages during summer, and that restingeggs facilitated population survival during the summer periodof stratification.     

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.     

Phytoplankton studies in the fjords of West Spitzbergen: physical environment and production in spring and summer

The phytoplankton in the fjords of West Spitzbergen was investigatedfrom 1978 to 1985. The spring bloom lasted longer than at theNorwegian coast: from mid March to early June. There is no delayin the onset of the spring bloom in Spitzerbergen fjords relativeto fjords of northern Norway. This is probably related to therapid daylength increase at high latitudes. The phytoplanktonspecies composition during spring was comparable to that alongthe coast of northern Norway. Annual primary production in theinvestigated area was calculated to be 150 g C m^–2 year^–1.     

Dinoflagellate bloom development and collapse in Lake Kinneret: a sediment trap study

Warm monomictic Lake Kinneret, Israel, is characterized by awinter–spring water bloom of the large (~50 µm diameter)dinoflagellate Peridinium gatunense Nygaard. Usually the P.gatunense bloom declines in May–June and a less prominentbloom of smaller dinoflagellates (mostly Peridiniopsis spp.of ~20–30 µm diameter) develops. Water column abundancesand sedimentation losses to those dinoflagellates were followedthroughout 1994 and 1995. The objective was to quantify thevariables that describe population dynamics, that in turn willshed more light on the seasonal patterns of bloom dynamics.Sedimentation losses were measured by means of sediment trapswith and without a preservative (formaldehyde) that were exposedfor 24 h once every 2–3 weeks. Annual sedimentation lossesof Peridinium (hypolimnetic trap catches) were 209 g wet wtm^-2 year^-1 in 1994 and 187 g wet wt m^-2 year^-1 in 1995, whichconstituted 16 and 23% of Peridinium production in those years,respectively. This study revealed that increased death ratespreceded a mass sedimentation flux of Peridinium and causedthe decline of the bloom in Lake Kinneret. Annual sedimentationlosses of Peridiniopsis were 55 g wet wt m^-2 year^-1 in 1994and 34 g wet wt m^-2 year^-1 in 1995. In contrast to live Peridiniumcells, Peridiniopsis cells continued to swim to the lower trapafter the onset of thermal stratification, possibly taking advantageof the higher nutrient concentrations below the thermocline,at a time when the lake is already stratified and the epilimnionis nutrient depleted. This could be an important factor allowingPeridiniopsis spp. to peak after the decline of Peridinium.     

Modelling of autumn plankton bloom dynamics

A simple system of parametrically forced ordinary differentialequations is used to model autumn phytoplankton blooms in temperateoceans by a mechanism involving deepening of the upper mixedlayer. Blooms are triggered provided the increase in nutrientsin the mixed layer is rapid within the first few days of deepeningand provided light-limited phytoplankton growth rate is relativelyhigh. Blooms exist as transient trajectories between quasi-equilibriumstates, rather than as bifurcations of steady states; thereforevery gradual deepening cannot trigger blooms. Very rapid deepeningalso prevents blooms due to the deleterious effect on phytoplanktongrowth rate. The mechanisms identified by this simple modelare vindicated by considering alternative grazing and deepeningregimes and by comparison with a more ecologically complex model(Fasham, 1993, in The Global Carbon Cycle, Springer-Verlag).Modelled estimates of primary productivity from both the simplemodel and the complex model parameterized for Ocean WeatherStation ‘India’ are around 0.5 g C m^–2 day^–1during the autumn bloom, therefore comprising a significantcomponent of annual production in temperate areas.     

Phytoplankton sinking rates in the Rhine region of freshwater influence

According to Stokes’ law, colony formation in phytoplanktonwould lead to enhanced sinking rates and higher sedimentationlosses if colonies had the same densities as the phytoplanktoncells they contain. In the Dutch coastal zone of the North Sea,algae settling out of the water column are subject to zoobenthosgrazing or to physical mixing into the sediment and, therefore,the formation of colonies by common diatom species and the prymnesiophytePhaeocystis globosa seems paradoxical: it would increase theprobability that sedimentation becomes a significant loss factor.However, sinking rate measurements in the Rhine region of freshwaterinfluence (ROFI) using SetCol settling columns did not reveala straightforward relationship between phytoplankton sizes (<10to >1000 µm) and sinking rates (–0.4 to >2.2m day^-1) of 24 autotrophic phytoplankton species and groups.In fact, under nutrient-replete conditions, the sinking ratesof the diatoms Chaetoceros radicans, Rhizosolenia shrubsoleiand Rhizosolenia stolterfothii decreased with size. The sinkingrates of large colonies of the prymnesiophyte P. globosa werealso negatively correlated with their size and positive buoyancywas observed. Chlorophyll a sinking rates exceeded 1 m day^-1periodically, which is sufficient to cause significant surfacelayer loss rates over 0.2 day^-1. Under stratified conditions,both chlorophyll a concentrations and sinking rates in the bottomlayer were significantly higher (+49% and +16%, respectively)than in the surface layer. These observations are discussedin relation to Stokes’ law, together with a critical analysisof the SetCol technique. It is concluded that: (i) SetCol givesadequate results when incubations are performed at or near insitu irradiance and temperature; (ii) sinking rates are predominantlydetermined by cell or colony density rather than their size;(iii) periodic sedimentation is an important species-specificloss process for phytoplankton in the Dutch coastal zone. Itis speculated that for diatoms with low sinking rates, autolysisis an important loss factor.     

Bloom of Emiliania huxleyi (Prymnesiophyceae) in the western South Atlantic Ocean

A monospecific bloom of the coccolithophorid Emiliania huxleyi(Lohmann) Hay & Mohler (Prymnesiophyceae) was detected offRio de La Plata at 36°S and 54°W in November 1989. Thecell densities observed were up to 6x10⁵ cells I^–1. Thisis the first record of a bloom of E.huxleyi in the area.     

Development of Anabaena blooms in a small reservoir with dense sediment akinete population, with special reference to temperature and irradiance

The development of Anabaena ucrainica blooms in a small agriculturalreservoir was monitored in 1998 and 1999. In the reservoir,numerous Anabaena akinetes were found in all regions of thesediment analyzed, with an average cell density in the uppermostlayer (0–2 cm) of 1.5 x 10⁴ cm^-3. Anabaena ucrainica filamentnumbers began to increase exponentially in mid-May 1998 andin late April 1999, when the water temperature exceeded 15°C.The average in situ net growth rate was 0.18 day^-1 as measuredby filament numbers. The effect of temperature on germinationof the akinetes was investigated using Anabaena akinetes takenfrom the reservoir sediment. High germination percentages wereobserved at temperatures between 14 and 23°C; however, theAnabaena akinetes did not germinate without irradiance. Growthexperiments using an axenic culture of A. ucrainica isolatedfrom the reservoir showed that an increase in incubation temperatureto 26°C resulted in a rise in the specific growth rate.Consequently, it was hypothesized that temperature increasescould similarly enhance the growth rate of A. ucrainica duringbloom development. Furthermore, judging from the in situ growthrate of A. ucrainica, initial inocula arising from dense akinetepopulations in the sediment would advance bloom formation andcould enhance the relative probability of Anabaena bloom formation.