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.     

Summer coastal zooplankton biomass and copepod community structure near the Italian Terra Nova Base (Terra Nova Bay, Ross Sea, Antarctica)

The structure of the zooplankton biotic community and of copepodpopulation in the coastal area of Terra Nova Bay (Ross Sea,Antarctica) was investigated during the 10th Italian AntarcticExpedition (1994/1995). Zooplankton biotic community consistedmainly of pteropods (Limacina helicina and Clione antarctica),Cyclopoid (Oithona similis), Poecilostomatoid (Oncaea curvata)and Calanoid (Ctenocalanus vanus, Paraeuchaeta antarctica, Metridiagerlachei and Stephos longipes) copepods, ostracods, larvalpolychaetes and larval euphausiids. Zooplankton abundance rangedfrom 48.1 ind m^–3 to 5968.9 ind m^–3, and copepodabundance ranged from 45.2 ind m^–3 to 3965.3 ind m^–3.The highest peak of zooplankton abundance was observed between25 m and the surface and was mainly due to the contributionof O. similis, O. curvata and C. vanus. Zooplankton biomassranged from 5.28 mg m^–3 to 13.04 mg m^–3 dry weight;the maximum value was observed between 25 m and the surface.Total lipid content varied from 216.44 to 460.73 mg g^–1dry weight.     

Biomass, feeding and production of Noctiluca scintillans in the Seto Inland Sea, Japan

The abundance and biomass of the large heterotrophic dinoflagellateNoctiluca scintillans, together with the changes in its potentialprey items, were monitored in the Seto Inland Sea, Japan, duringsummer 1997 (17 July-11 August). Growth and grazing rates ofNscintillans fed natural plankton populations were also measuredeight and seven times, respectively, during the survey period.The abundance and biomass of N scintillans averaged over thewater column (19 m) were in the range 1–345 cells 1^–1(temporalaverage = 93 cell1^–1) and 0.1–49.6 µg C l^–1(temporalaverage = 13.8 µg C l^–1; three times higher thanthat of calanoid copepods during the same period). Noctilucascintillans populations followed the changes in phytoplankton:N.scintillans biomass was increasing during the period of diatomblooms and was at a plateau or decreasing during periods oflow chlorophyll a. The growth rates of N.scintillans (µ)were also consistent with the wax and wane of the N.scintillanspopulation: N.scintillans showed highest growth rates duringdiatom blooms. A simple relationship between µ and chlorophylla concentration was established, and the production of N.scintillanswas estimated using this relationship and the measured biomass.The estimated production averaged over the water column wasin the range >0.1–5.2 µg C l^–1 day^–1(temporalaverage = 1.4 µg C l^–1 day^–1; 64% of the productionof calanoid copepods during the same period). Diatom clearancerates by N.scintillans were in the range 0.10–0.35 mlcell^–1 day^–1, and the phytoplankton population clearanceby N.scintillans was >12% day^–1. Thus, although thefeeding pressure of N.scintillans on phytoplankton standingstock was low, N.scintillans was an important member of themesozooplank-ton in terms of biomass and production in the SetoInland Sea during summer.     

Biomass and size structure of the scyphomedusa Aurelia aurita in the northwestern Black Sea during spring and summer

The abundance, biomass and size structure of the scyphomedusa,Aurelia aurita, was measured during two research cruises tothe northwestern Black Sea (July–August 1995 and April–May1997). Average biomass of Aurelia was relatively constant (132–179g wwt m^–2) throughout the investigation period and similarto previous years. Abundance and biomass at individual stationsappeared to be unrelated to temperature and salinity when thelatter exceeded ~13. Biomass was low at coastal stations inthe plume of the Danube where depth was <20 m and salinitydropped to <11. The spring cruise (April–May) coincidedwith, or just followed the peak of strobilation. The summercruise (July–August) took place near the beginning ofplanulae larvae release. The population size structure was dominatedby small individuals in spring, while large medusae prevailedmainly in late summer. Aurelia was, on average, larger at deepwater stations during summer, suggesting that per capita foodsupply was higher further offshore. The individual body massincreased from spring through summer. Accordingly, the volume(wet wt) to length (bell diameter) relation changed significantly.If all medusae measured throughout the seasons were pooled,volume (V, in cm³) was related to length (L, in cm) accordingto V = 0.08 L^2.71, which is similar to measurements conductedin other coastal areas. In contrast to the common conjecture,we did not find inverse relations between biomasses of Aureliaand the combjelly Mnemiopsis leidyi. Preliminary feeding experimentsindicate that Aurelia may feed upon small Mnemiopsis. The significanceof indirect trophic relations and direct feeding interactionsamong the gelatinous zooplankton in the Black Sea has importantconsequences for the energy flow along the food web and, therefore,needs further study.     

Gymnodinium catenatum Graham (Dinophyceae)in Europe: a growing problem?

The microreticulate resting cyst of the potentially toxic, chain-forming,unarmoured neritic dinoflagellate Gymnodinium catenalum Graham1943. the planktonic stage of which is not known from NorthEuropean waters, is reported for the first time from recentGerman coastal sediments of the North Sea and Baltic Sea. Insandy mud sediments of the German Bight, a maximum of 8 5 livingcysts cm^–3 were found. In Kiel Bight sediments G.catenalumwas found in maximum concentrations of 17.0 living cysts cm^–3.In surface waters of the German Bight resuspended G catenatumcysts were observed at concentrations of up to 3.6 cysts l^–1.Successful germination experiments conducted with natural seawatershow that the occurrence of a vegetative form of G.catenatumin northern Europe is very likely. The present study highlightsthat cyst surveys provide an important tool for the evaluationof areas with potential toxicity problems, as they may indicatethe presence of hitherto overlooked species in the water column.     

Summer phytoplankton succession in Ellis Fjord, eastern Antarctica

Spring phytoplankton communities in the water column of EllisFjord are characterized by diatoms originating from the bottomsea-ice strand community. Upon ice breakout in early summer,these are replaced by blooms of the phytoflagellates, Phaeocystispouchetii, Cryptomonas cryophila, Pyramimonas gelidicola, silicoflagellatcsand dinoflagellates. The narrow entrance of the fjord and thedevelopment of summer stratification is probably limiting theavailability of nutrients and containing the magnitude of thesmall bloom (maximum 2.8 10⁶ cells l^–1).     

Effect of light regime upon growth rate and chemical composition of a clone of Skeletonema costatum from the Trondheimsfjord, Norway

Nutrient-saturated cultures of Skeletonema costatum were grownat 15C and 42 combinations of photon flux density (PFD) anddaylength. The growth rate increased with the daylength andPFD up to 460–630 µE m^–2 s^–1 (maximum2.5 doublings day At 2000 µE m^–2 s^–1 the growthrate was reduced by 45%. The chlorophyll (chl) content of thecells and the rate of production of carbon per unit chlorophylland ambient light increased for declining light regimes as didcellular nitrogen and carbon. The N/C ratio, cellular phosphorusand ratios between in vivo fluorescence, with and without DCMU,and chlorophyll varied negligibly. The ATP/C ratio was linearlyrelated to the growth rate. The results were described mathematically.The chl/C ratio was low both in strong light and in marginallylow light, corresponding to low cellular chlorophyll and highcellular carbon, respectively. The observed increase in cellularnitrogen and carbon at shade adaptation probably represent anincrease in the size of internal stores of organic nitrogenand may imply that Skeletonema cells become enriched with organicnitrogen when staying in nitrate-rich subsurface layers, e.g.in or below a nutricline. However, close to zero growth in marginallight the cells become greatly enriched with respect to everymeasured factor. Such cells may be physiologically resting stageswhich may ensure survival during dark periods and promote rapiddevelopment during the initial phase of blooms. Cultures andnatural blooms of Skeleronema in the Trondheimsfjord exhibitvery similar patterns of variation.     

The contribution of ammonia excreted by zooplankton to phytoplankton production in Narragansett Bay

Ammonia excreted by mixed zooplankton populations over an annual(1972–1973) cycle in Narragansett Bay varied from 0.04to 3.21 µg at NH₃-N dry wt^–1 day^–1, exclusiveof two exceptional rates measured one year apart: 11.74 and18.39 µg at NH³-N mg dry wt^–1 day^–1. Grossphytoplankton production integrated over the year (1972–1973)averaged 151 mg C m^–3 day^–1 for an 8 m water column;peaks of 332 and 905 mg C m^–3 day^–1 occurred duringthe winter-spring and summer blooms, respectively. Excretedammonia, integrated seasonally and annually, contributed only0.2% and 4.9% of the nitrogen required for observed gross productionduring the winter-spring and summer blooms, respectively, and4.4% annually. However, excreted ammonia may be an importantsource of the nitrogen required by Skeletonema costatum, thedominant diatom in Narragansett Bay, during the post-bloom periodwhen 186% of the nitrogen required for its net production wasmet by ammonia excretion. A combination of zooplankton ammoniaexcretion and benthic ammonia flux contributed 22% of the nitrogenrequired for the annual gross production (440 g C m^–2)while 51% of the nitrogen required for the net production ofSkeletonema was accounted for by regenerated nitrogen. ¹This research was supported by NSF grant GA 31319X awardedto Dr.T.J.Smayda.     

Size-fractionated biomass, photosynthesis and dark CO2 fixation in a tropical oceanic environment

This study examines the spatial distribution and size structureof phytoplankton biomass and productivity in relation to thevertical structrure of the Andaman Sea (northeastern IndianOcean). This region was characterized by low concentrationsof nutrients and high levels of insolation. Nitrogen availabilityappeared to control overall productivity with nitrate-based‘new’ production accounting for 8–24% of thetotal primary production. Euphotic column chlorophyll (chl a)averaged 52.5 mg m^–2 of which a major portion was locatedas a subsurface chl a maximum (SCM) at     

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.     

Plankton community respiration in Bransfield Strait (Antarctic Ocean) during austral spring

Community respiration (R) was determined in Bransfield Straitfrom oxygen changes in water samples incubated in borosilicatebottles maintained at in situ temperature. The respiratory electrontransport system (ETS) activity of seawater communities wasalso measured from the same samples. Both data sets were relatedby the regression equation: log R (mg O₂ m^–3 day^–1)=0.462+0.730xlogETS activity mg O₂ m^–3 day^–1) (r=0.80, n=23). Fromthis equation and 37 ETS activity depth profiles, we calculatedthe integrated (0–100 m) community respiration as beingin the range 1.2–4.5 g O₂ m^–2 day^–1 (mean=2.2).These values do not differ significantly from other publishedresults for the Arctic and Antarctic Oceans. Assuming a respiratoryquotient of unity, the areal respiration ranges between 0.45and 1.69 g C m^–2 day^–1 (mean=0.8). This would representan important sink for the primary production reported for BransStrait. The spatial distribution of community respiration showedhigher values associated with the warmer and phytoplankton-richwaters outflowing from Gerlache Strait into Bransfield Strait,and with the front that separates Bellingshausen Sea watersfrom Weddell Sea waters. We suggest that this pattern of distributionmay be related to the transport of organic matter by the BransfieldCurrent along the front.     

The productive and optical status of the oligotrophic waters of the Southern Aegean Sea (Cretan Sea), Eastern Mediterranean

Primary production, pigment concentrations and spectral measurementsof downwelling irradiance were made at four stations in fourseasons (spring, summer, autumn, winter) during 1994 in thewaters of the South Aegean Sea (Cretan Sea), Eastern Mediterranean.Rates of production were determined using in Situ incubationtechniques and included measurements at the surface microlayer.Depth-integrated values averaged over season were 5.66 mg Cm^–2 h^–1 for primary production and the correspondingchlorophyll (Ch1) a and phaeophytin (Phaeo) a values had meansof 4.87 and 1.21 mg m^–3 respectively. The assimilationratio remained very low (mean over season: 1.19 mg C mg^–2Chl a h^–1 as did the Phaeo a/Chl a ratio (mean over season:0.24). The annual production for the area was estimated to yield24.79 g C m^–2 year^–1. Primary production and Chla estimates showed statistically significant seasonal, spatialand depth variations. The spectral values of the attenuationcoefficient Kd (     

Zooplankton community structure and copepod egg production in coastal waters of the central Great Barrier Reef lagoon

We describe zooplankton community structure and copepod eggproduction in the vicinity of the coastal boundary zone of theGreat Barrier Reef lagoon, Australia. The abundance and eggproduction rate of constituents of the zooplankton assemblagecharacteristic of the coastal zone rapidly increase subsequentto events such as flooding and upwelling. Our sampling spannedtwo summer monsoonal seasons, the first of which (1990-91) wasvery wet. The second monsoonal season (1991-92) was very dryand was characterized by intrusive upwelling events from theCoral Sea. Chlorophyll a concentrations did not rise in thewet year, probably because of light limitation, but did riseas a result of upwelling. Terrestrial run-off in the wet yearhad a greater apparent effect on zooplankton abundance patternsthan did upwelling in the dry year, except where coastal trappingallowed sufficient time for increases in zooplankton abundanceto occur. Egg production rates by the copepods Acrocalanus gibberand Acrocalanus gracilis showed haphazard spatial differences.Nitrogen-specific egg production ranged between 0.03 and 0.21day^–1 for A.gibber, and between 0.13 and 0.41 day^–1for A.gracilis. The egg production rate by A.gibber was foodlimited for most of the year and showed a poor correlation withtemperature. ³Present address: Department of Biological Sciences, FloridaTech, 150 W University Boulevard, Melbourne, FL 32901, USA     

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.     

Discontinuite hydrobiologique dans le detroit du Pas-de-Calais

Hydrobiological studies in the Straits of Dover show a differencebetween the characteristics of the French coastal water andthe open sea water. A cruise between Boulogne-sur-mer and Dover(October 15, 1985) confirms the existence of a frontal area(4 miles off the French coast). This area is characterized byan increase of the salinity (1%) and decrease of the suspendedmatter content (from 27 to 20 mg l^–1), of primary production(from 15 to 3 mg C m^–3h^–1) and of chlorophyll aconcentration (from 4.5 to 1.5 mg m^–3). The presence ofdifferent zooplanktonic species in the two water masses alsocharacterizes this discontinuity. Oncaea sp., Corycaeus anglicus,Centropages typicus and Calanus helgolandicus are indicatorspecies of open sea water, while Cyciopina littoralis and ‘Saphirella’are only present in coastal water.     

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.     

Estimation of ammonium regeneration efficiencies associated with bacterivory in pelagic food webs via a 15N tracer method

Phagotrophic protists are major components of pelagic food webs,both as consumers of bacterial and phytoplankton cells, andas regenerators of inorganic nutrients. In this study, we estimatedthe efficiency of ammonium regeneration by protists feedingon bacteria within natural plank-tonic assemblages, using a¹⁵N tracer method, in which the excretion of ¹⁵N-labeled ammoniumdue to grazing on ¹⁵N pre-labeled bacteria was followed overtime. We tested this approach in experiments based on the additionof heat-killed ¹⁵N-labeled bacteria to laboratory cultures andto samples of coastal seawater. During two experiments, variationin abundance of bacterivores and bacterioplankton resulted innon-constant grazing rates. Deterministic computer models thatused abundance of bacteria and protists as variables were developedto estimate best-fit values of grazing mortality (g, h^–1)and of ammonium regeneration efficiency (R_E, fraction of theinitial ¹⁵N label in added bacteria which is released as ammonium).Estimated ammonium R_E were 0.30–0.35 for one trophic linksystems with both a monospecific culture and a mixed speciesassemblage of bacterivorous flagellates. R_E was higher for multi-trophicstep food webs: 0.60 for 5 µm pre-screened coastal seawaterand 0.90 for whole coastal seawater.     

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     

Springtime microprotozoan abundance and biomass in the southeastern Bering Sea and Shelik of Strait, Alaska

We surveyed springtime biomass and abundance of the >20 µmmicroprotozoa in surface waters of the SE Bering Sea and Shelikof Strait, Alaska. This study was part of the Fisheries OceanographyCoordinated Investigations (FOCI) program examining processeswhich affect the recruitment variability of walleye pollock(Theragra chalcogramma). Microprotozoa are a potential preyresource for larval pollock which has not been previously examined.In both areas, the >20 µm microprotozoa were predominantlydinoflagellates and ciliates. At the time of sampling (May 1990in Shelikof Strait and April 1992 in the SE Bering Sea), thespring diatom bloom was under way in Shelik of Strait, but notin the SE Bering Sea. Heterotrophic dinoflagellates dominatedthe microprotozoan assemblage in Shelik of Strait, but not inthe SE Bering Sea. In the SE Bering Sea. total microprotozoanabundances ranged from 300 to 6233 organisms 1^–1 and biomassfrom 0.58 to 9.73 µg C 1^–1. In Shelik of Strait,abundance and biomass were higher, ranging from 850 to 14 960organisms 1^–1 and from 1.29 to 70.73 µg C 1^–1,respectively. These biomass levels are comparable to those reportedfrom other coastal and oceanic regions. Microprotozoan biomasslevels were sufficient to support the estimated metabolic needsof first-feeding larval walleye pollock. It remains to be shownwhether larval pollock use this resource.     

Planktonic primary production in the German Wadden Sea

By combining weekly data of irradiance, attenuation and chlorophylla concentrations with photosynthesis (P) versus light intensity(E) curve characteristics, the annual cycle of planktonic primaryproduction in the estuarine part of the Northfrisian WaddenSea was computed for a 2 year period. Daily water column particulategross production ranged from 5 to 2200 mg C m^–2 day^–1and showed a seasonal pattern similar to chlorophyll a. Budgetcalculation yielded annual gross particulate primary productionsof 124 and 176 g C m^–2 year^–1 in 1995 and 1996,respectively. Annual amounts of phytoplankton respiration, calculatedaccording to a two-compartment model of Langdon [in Li,W.K.W.and Maestrini,S.Y. (eds), Measurement of Primary Productionfrom the Molecular to the Global Scale. International Councilfor the Exploration of the Sea, Copenhagen, 1993, pp. 20–36],and dissolved production in 1996, were both in the range of24–39 g C m^–2 year^–1. Annual total net productionwas thus very similar to particulate gross production (127 and177 g C m^–2 year^–1 in 1995 and 1996, respectively).Phytoplankton growth was low or even negative in winter. Inspring and summer, production/biomass (Pr/B) ratios varied from0.2 up to 1.7. Phytoplankton growth during the growth seasonalways surpassed average flushing time in the area, thus underliningthe potential of local phytoplankton bloom development in thispart of the Wadden Sea. The chlorophyll-specific maximum photosyntheticrate (P^B_max) ranged from 0.8 to 9.9 mg C mg^–1 Chl h^–1and was strongly correlated with water temperature (r² = 0.67).By contrast, there was no clear seasonal cycle in ^B, which rangedfrom 0.007 to 0.039 mg C mg^–1 Chl h^–1 (µmolphotons m^–2 s^–1)^–1. Its variability was muchless than P^B_max and independent of temperature. The magnitudeand part of the variability of P^B_max and ^B are presumably causedby changes in species composition, as evidenced from the rangeof these parameters found among 10 predominant diatom speciesisolated from the Wadden Sea. The ratio of average light conditionsin the water column (E_av) to the light saturation parameterE_k indicates that primary production in the Wadden Sea regionunder study is predominantly controlled by light limitationand that nutrient limitation was likely to occur for a few hoursper day only during 5 (dissolved inorganic nitrogen) to 10 (PO₄,Si) weeks in the 2 year period investigated.