Colonization of biofilms by bacteria capable of biodegrading sodium dodecyl sulphate (SDS) at clean and polluted riverine sites

Fifty cyanobacterial strains (10 genera) were tested in batch culture for their ability to use organic phosphorus compounds (1 mg liter⁻¹ P) as their sole P source. Two monoesters, Na₂-β-glycerophosphate and π-nitrophenyl phosphate (πNPP), supported growth of all strains, and the diester bis-π-nitrophenyl phosphate (bis-π-NPP) and herring sperm DNA supported almost all strains. ATP was either a very favorable or poor P source and failed to support growth of nine strains, seven of which were Rivulariaceae with trichomes ending in a hair or long tapered region. Phytic acid was in general the least favorable P source. P-limited cultures grown initially with inorganic phosphate to conditions of P limitation were also tested for cell-bound and extracellular phosphomonoesterase (PMEase) and phosphodiesterase (PDEase) activities at two pH values (7.6, 10.3) using πNPP and bis-πNPP as substrates. Cell-bound PMEase was inducible in all strains and cell-bound PDEase in most strains. Most showed extracellular PMEase, but not extracellular PDEase. The highest values (μM πNPP or bis-πNPP hydrolyzed mg dry weight⁻¹ hour⁻¹) all occurred in strains ofGloeotrichia as follows: cell-bound PMEase at pH 7.6, 2.7 μM in strain D602; cell-bound PMEase at pH 10.3, 5.2 μM in D602; extracellular PMEase at pH 7.6, 0.73 μM in D281; extracellular PMEase at pH 10.3, 6.6 μM in D281; cell-bound PDEase at 7.6, 0.40 μM in D613; cell-bound PDEase at pH 10.3, 1.0 μM in D613. The results were compared to see if they indicated possible relationships between phosphatase activity and taxonomic or ecological grouping. The following differences were significant (P<0.05). Rivulariaceae produced higher yields than filamentous non-Rivulariaceae with β-glycerophosphate, πNPP, and DNA. Rivulariaceae with the ability to form hairs in culture showed poorer growth in ATP than non-hair-forming Rivulariaceae, but were more effective at utilizing phytic acid. Strains from calcareous environments had higher PMEase activity at pH 10.3 than strains from noncalcareous environments (P<0.01).     

Phosphatase activities of the aquatic moss <Emphasis Type="Italic">Warnstorfia fluitans</Emphasis> (Hedw.) Loeske from an acidic stream in North-East England

A study was made of the aquatic environment, tissue nutrient composition and surface phosphatase activities of the aquatic moss Warnstorfia fluitans in Brandon Pithouse Stream, a small acidic stream in N-E England. The water, which originates from an underground spring, had been pH 2.6 for at least 30 years, but about 3.9 during the present study. The moss was by far the most abundant phototroph during all this period. Seasonal changes in aqueous nitrogen and phosphorus fractions were measured over a 2-year period near the source. Most of the filtrable N and P were at times organic, but the very high N:P ratio (even if organic N is excluded) suggests that only organic phosphate is likely to be important for the moss. There was a high peak in organic phosphate in late spring in both study years. Surface phosphomonoesterase (PMEase) and phosphodiesterase (PDEase) activities were highly correlated in the field and in axenic culture, though there were some differences in response to environmental factors. Axenic material showed higher PMEase and PDEase activities when grown with organic P than with inorganic P. Although the data suggest that internal P content is an important factor influencing phosphatase activities, PDEase activity was especially marked when the moss was grown with the diester, DNA, as P source, indicating that at least one of its surface phosphatases can also respond directly to the environment. Handling editor: S. M. Thomaz     

Preferential utilization of inorganic polyphosphate over other bioavailable phosphorus sources by the model diatoms Thalassiosira spp.

Polyphosphates and phosphomonoesters are dominant components of marine dissolved organic phosphorus (DOP). Collectively, DOP represents an important nutritional phosphorus (P) source for phytoplankton growth in the ocean, but the contribution of specific DOP sources to microbial community P demand is not fully understood. In a prior study, it was reported that inorganic polyphosphate was not bioavailable to the model diatoms Thalassiosira weissflogii and Thalassiosira pseudonana. However, in this study, we show that the previous finding was a misinterpretation based on a technical artefact of media preparation and that inorganic polyphosphate is actually widely bioavailable to Thalassiosira spp. In fact, orthophosphate, inorganic tripolyphosphate (3polyP), adenosine triphosphate (ATP) and adenosine monophosphate supported equivalent growth rates and final growth yields within each of four strains of Thalassiosira spp. However, enzyme activity assays revealed in all cultures that cell-associated hydrolysis rates of 3polyP were typically more than ~10-fold higher than degradation of ATP and the model phosphomonoester compound 4-methylumbelliferyl phosphate. These results build on prior work, which showed the preferential utilization of polyphosphates in the cell-free exudates of Thalassiosira spp., and suggest that inorganic polyphosphates may be a key bioavailable source of P for marine phytoplankton.     

Weak 'top-down' control of dinoflagellate growth in the coastal Skagerrak

Effects of planktivorous fish (Sprattus sprartus), copepods(mainly Oithona sp.) and benthic filter feeders (Mytilus edulis)on the biomass and species composition of a natural marine planktoncommunity, dominated by dinoflagellates, was studied experimentallyin September 1988. Plastic 300 I cylinders were filled withSkagerrak surface water, filtered through a 90 µm nylonnet in order to remove mesozooplankton. Copepods were addedin three different concentrations (0, 1x and 6x the ambientdensity) with and without nutrient additions (nitrate, silicateand phosphate). In two nutrient-enriched cylinders with addedcopepods (1x and 6x), 5 cm sprats were also added (10 per cylinder).In two nutrient-enriched cylinders without added copepods. bluemussels (20 per cylinder. 3 cm length) were introduced. Chlorophylla. inorganic nutrients and densities of phytoplankton, ciliatesand copepods were measured throughout the experimental periodof 10 days. Gyrodinium aureolum increased in cell numbers inall but the mussel cylinders, while the other dominating dinoflagellate,Ceratium furca, decreased. This decrease was not due to grazing,but probably a container artefact. Diatoms, Rhizosolenia fragilissinia,Leptocylindrus danicus and Chaetoceros spp., increased slowlyin treatments without fish. In the fish treatments, diatomshad appreciably higher growth rates and cell numbers, probablydue to predation by fish on the copepods. Gyrodiniun aureolunalso had a somewhat higher growth rate in the presence of fish.although this was probably not due to reduction of copepod grazing.The results show that planktivorous fish can affect the biomassand species composition of natural phytoplankton communitiesin marine waters, but that the effect on the late summer-autumndinoflagellate dominated community is weak. Blue mussels areefficient grazers of the whole phytoplankton community and arenot particularly selective with respect to either size or species.     

Changes in photosynthetic activity of the diatom Phaeodactylum tricornutum in a culture of limited volume

Changes in photosynthetic activity of a marine diatom duringalgal growth were studied with a typical culture medium formarine algae, ASP-2 (l). As the algal specimen, Phaeodactylumtricornutum was used for the experiments. Nitzschia closteriumand Chaetoceros sp. were also supplementarily used. Photosynthetic and p-benzoquinone Hill activities remarkablychanged with time during algalgrowth; with maximum activityfound in cells at log phase. A rapid decrease occurred in theinterphase from the log to stationary phase. The activity changewas not accompanied by variation in photosynthetic pigment content. The low concentration of phosphorus source was suggested asthe main cause for the change. On supply of extra inorganicphosphate, the time length for holding high photosynthetic activitybecame longer; or, the activity of the cells at stationary phaserecovered at least partly even in the dark. Dark recovery wasnot accompanied by either algal growth or an increase in thecontent of photosynthetic pigments. Inactivation of photosynthesis in the stationary growth phaseand activation by added phosphate in the dark were inferredto be due to changes in activities of both the CO₂-fixing andelectron transfer systems. The observed activity change maynot be attributable to a deficiency in inorganic phosphate asthe substrate for photophosphorylation. Similar changes in photosynthetic activity were also observedwith Nitzschia closterium and Chaetoceros sp. (Received January 30, 1971; )     

Do transparent exopolymer particles (TEP) inhibit grazing by the euphausiid Euphausia pacifica?

The hypothesis that ubiquitous, sticky transparent exopolymerparticles (TEP) formed from phytoplankton exudates will adhereto and coat the feeding structures of marine zooplankton grazers,and thus depress feeding on phytoplankton, was tested usingthe euphausiid, Euphausia pacifica, as a model organism. Duringtwo feeding experiments, E.pacifica were offered cells of thediatom Thalassiosira weissflogii, TEP, or both TEP and T.weissflogiicells. Ingestion rates on cells were lower in the presence ofTEP. However, contrary to the hypothesis, grazing on cells wasnot inhibited by TEP. Rather, TEP-clusters, aggregates whichformed from TEP and nano-sized particles normally too smallfor the filtering apparatus of E.pacifica to retain, servedas an alternative food source for E.pacifica, reducing theiringestion of cells. These clusters were very similar in formto the TEP actually available to marine grazers in nature. TEP-clusterswere similar to cells in size and food quality, and were grazedat similar rates. When feeding on TEP-clusters, euphausiidsshort circuit the food web by feeding on nano- and picoplanktondirectly, bypassing the microbial loop. Thus, the presence ofTEP appears to enhance, rather than depress, macrozooplanktongrazing.     

Advantages from diel vertical migration can explain the dominance of Gonyostomum semen (Raphidophyceae) in a small, steeply-stratified humic lake

In late summer, a large flagellated alga, Gonyostomum semen(Raphidophyceae), constituted most of the phytoplankton biomassin a small steeply-stratified humic lake. Its diel verticalmigration (DVM) was very distinct and extended at night intothe anoxic hypolimnion. After midsummer, the depletion of hypolimneticoxygen led to a gradual release of soluble reactive phosphorus(SRP) from the sediment, but one month later, irrespective ofcontinuing stratification, the concentrations again returnedto undetectable levels down to the bottom. As this coincidedwith the rapid increase in G.semen population, the latter wasprobably responsible for the depletion of SRP. The flux of SRPfrom the sediment to the epilimnion was virtually interrupted,making non-migrating phytoplankton dependent only on regeneratedand inflowing inorganic phosphorus. Besides nutrient availability,DVM also benefited G.semen in the reduction of metabolic andgrazing losses. In this lake, the remarkable multiple advantagesof DVM probably explain the dominance of the large G.semen inthe late summer phytoplankton biomass over much smaller algae.     

Phytoplankton and periphyton responses to in situ experimental nutrient enrichment in a shallow subtropical lake

Experiments were performed in situ in shallow, subtropical LakeOkeechobee (Florida. USA) to quantify and compare the responsesof phytoplanklon (in 20 I clear polycarbonate carboys) and periphyton(on nutrient-diffusing clay substrates) to additions of nitrogenand/or phosphorus. During early and late summer. 1994, bothassemblages were nitrogen limited or co-limited by nitrogenand phosphorus, indicating the potential for competition betweenbenthic and planktonic communities. During late summer, therewas evidence that high phytoplankton biomass reduced light penetrationthrough the water column and may have suppressed periphytongrowth. The similar phytoplankton and periphyton taxonomic structures,both dominated by Lyngbya sp. and pennate diatoms, suggestedthat in shallow regions of this lake, resuspended meroplanktonmight account for a large portion of phytoplankton biomass.This phenomenon has been observed in other shallow, wind-drivenFlorida lakes.     

Alkaline phosphatase activities of anAnabaena sp. from deep-water rice

Anabaena sp. grew with mono- and di-ester phosphate compounds as sources of phosphate, indicating the presence of phosphomonoesterase (PMEase) and phosphodiesterase (PDEase) activities. Cell-bound PMEase and PDEase activities were detected during growth in 0.5 and 10 mg PO₄l^–1 only when the cellular phosphate concentration fell to 0.46% of cell protein and the activities increased as cellular phosphate content decreased. The K_m values for these enzymes were 0.3mm forp-nitrophenyl phosphate and 0.2mm for bis-p-nitrophenyl phosphate, respectively. Only PMEase activity was found extracellularly. The pH optima for PMEase and PDEase were 10.2 and 10.4, respectively, and the temperature optima at pH 10.2 were 37°C and 40°C, respectively. Ca²⁺ increased the enzyme activities while Zn²⁺ caused marked inhibition. The inorganic phosphate repressed the cellular PMEase activity after a lag of 4 h.     

Experimental evaluation of herbivory in the ctenophore Mnemiopsis leidyi relevant to ctenophore-zooplankton-phytoplankton interactions in Narragansett Bay, Rhode Island, USA

Herbivory of Mnemiopsis leidyi and its interactions with phytoplanktonand non-gelatinous zooplankton were examined in small-scalemicrocosm experiments. Clearance rates for M. leidyi incubatedwith phytoplankton were generally negative, but ranged up to4.5 1 ctenophore^–1 day^–1 when the large (80 µmø) diatom Ditylum brightwelli was offered as a food source.These highest ingestion rates would provide Mnemiopsis withonly 21 % of its daily carbon requirements for respiration.Mean shrinkage of M. leidyi was 8.2–51% when incubatedwith phytoplankton. Although M. leidyi neither fed activelyon phytoplankton, nor satisfied its nutritional needs on sucha diet, the chain-forming diatom Skeletonema costatum becameentangled in mucus strands and balls produced by M. leidyi inthe absence of zooplankton. Attachment onto mucus occurred atphytoplankton concentrations commonly observed in NarragansettBay and may be important in the formation of "marine snow" duringsummer M. leidyi pulses; phytoplankton sinking rate and the"package size" available to herbivores would also be affected.The experiments support our previous hypothesis based on fieldobservations in Narragansett Bay that M. leidyi indirectly regulatesphytoplankton abundance there during the summer bloom as a consequenceof predation on zooplankton. The extent to which M. leidyi influencedphytoplankton dynamics in the microcosms was dependent on therelative abundance and physiological state of the three trophiclevels. A food web diagram for M. leidyi is presented.     

Enclosure Experiments with Low-dose Additions of Phosphorus and Nitrogen in the Acidified Lake Njupfatet,Central Sweden

Experiments involving low-dose additions of phosphate, ammonium, nitrate and ADP, one by one and in combination, were performed in small (350 litre) in-situ enclosures in a moderately acid (pH 5.4) lake. Before manipulation, all large filter-feeding animals were removed by filtration. Phytoplankton responded to the nutrient additions only when both phosphorus and nitrogen were added, thus indicating a close balance between phosphorus and nitrogen limitation in the system. Variation of the inorganic nitrogen-source resulted in species-specific responses by phytoplankton. With ammonium as the nitrogen source Merismopedia tenuissima was favoured, regardless of whether this species was dominant in the phytoplankton community at the beginning of the experiment or not. With nitrate as nitrogen source Peridinium inconspicuum, which was never particularly common at the beginning of the experiments, was favoured. No other species of phytoplankton present in the bags was able to outcompete these two species as long as inorganic nutrients were added. With ADP as phosphorus source together with nitrate, a third species, Dictyosphaerium cf. botrytella, was favoured and reached dominance. The zooplankton community remaining in the bags, dominated by rotifers and calanoid nauplii, did not respond to the fertilization-induced increases in the total biomass of phytoplankton.     

Phytoplankton and zooplankton development in a lowland, temperate river

The longitudinal and seasonal patterns of plankton developmentwere examined over 2 years in a lowland, temperate river: theRideau River (Ontario, Canada). Following an initial decreasein phytoplankton and zooplankton biomass as water flowed fromthe headwaters into the Rideau River proper, there was an increasein chlorophyll a (chl a) and zooplankton biomass with downstreamtravel. At approximately river km 60, both phytoplankton andzooplankton reached their maximum biomass of 27 µg l^–1(chl a) and 470 µg l^–1 (dry mass), respectively.Downstream of river km 60, the biomass of both planktonic communitiesdeclined significantly despite increasing nutrient concentrationsand favorable light conditions. These downstream declines maybe due to the feeding activity of the exotic zebra mussel (Dreissenapolymorpha) which was at high density in downstream reaches(>1000 individuals m^–2). There was no evidence forlongitudinal phasing of phytoplankton and zooplankton, as increasesand decreases in chl a and zooplankton biomass appeared to coincide.Overall, chl a was best predicted by total phosphorus (R²=0.43),whereas zooplankton biomass was best predicted by chl a (R²=0.20).There was no evidence for significant grazing effects of zooplanktonon phytoplankton biomass.     

Phytoplankton seasonal dynamics in a Mediterranean coastal lagoon: emphasis on the picoeukaryote community

The dynamics of the phytoplankton community were investigatedin a marine coastal lagoon (Thau, NW Mediterranean) from February1999 to January 2000. Dilution experiments, chlorophyll a (Chla) size-fractionation and primary production measurements wereconducted monthly. Maximum growth and microzooplankton grazingrates were estimated from Chl a biomass fractions to separatepico- from nano- and microphytoplankton and by flow cytometryto distinguish between picoeukaryotes and picocyanobacteria.In spring, the phytoplankton community was dominated by Chaetocerossp. and Skeletonema costatum, which represented most of biomass(B) and primary production (P). Nano- and microphytoplanktongrowth was controlled by nutrient availability and exceededlosses due to microzooplankton grazing (g). Picoeukaryote andcyanobacteria growth was positively correlated with water temperatureand/or irradiance, reaching maximum values in the summer (2.38and 1.44 day^–1 for picoeukaryotes and cyanobacteria, respectively).Picophytoplankton accounted for 57% of the biomass-specificprimary productivity (P/B). Picophytoplankton was strongly controlledby protist grazers (g = 0.09–1.66 day^–1 for picoeukaryotes,g = 0.25–1.17 day^–1 for cyanobacteria), and microzooplanktonconsumption removed 71% of the daily picoplanktonic growth.Picoeukaryotes, which numerically dominate the picoplanktoncommunity, are an important source of organic carbon for theprotistan community and contribute to the carbon flow to highertrophic levels.     

Phosphorus utilization by Asterionella formosa Hass

Observations have been made on the relationship in Lake Windermerebetween the growth of Asterionella formosa and the concentrationof dissolved phosphate. Asterionellaformosa has also been grownin culture and the amount of phosphorus required by this organismhas been determined. These experiments have shown that: (1) Asterionella can take up and store in reserve phosphorusfrom concentrations below those found in phosphorus-poor lakes(i.e. below 1 µg.P/l.). (2) Growth can continue in phosphorus-deficient media by makinguse of such reserves, cell phosphorus being steadily reduced. (3) The limiting requirement per cell of phosphorus is veryminute—about 0—06µg P/10⁶ cells/1, so thatinitial concentrations as low as rog.P/l. can theoreticallyproduce a population of some i6 x io6 cells/l. before limitationby phosphorus deficiency. This has been realized in culture.The behaviour of Asterionella formosa growing in nature hasbeen found to conform with that found in culture. It is concludedfrom such observations that phosphorus deficiency is unlikelyto provide a limit to growth of Asterionella in Windermere,despite the very low initial concentration of dissolved phosphate.Further experiments have shown that Asterionella cells low inphosphorus can rapidly take up added phosphate from lake waterbut not from distilled water. Some factors which affect therate of phosphate uptake of depleted cells are investigated,and attempts have ben made to throw some light on the natureof the apparent difference between lake water on the one handand distilled water and a number of artificial lake waters onthe other. No conclusion is reached on the reason why Asterionellacells behave differently in lake water and in artificial media.     

Nitrogen and Phosphorus in a Stretch of the Guadalupe River,Texas, with Five Main-Stream Impoundments by

Nitrogen and phosphorus were studied in a 168-km stretch of the Guadalupe River that had five main-stream impoundments. Flow through the study area was controlled by releases from these five reservoirs and from Canyon Reservoir, a deep-storage reservoir, located 30 km upstream. Parameters measured monthly on a diel basis at 16 stations were nitrate nitrogen, nitrite nitrogen, ammonia nitrogen, Kjeldahl nitrogen, inorganic phosphate phosphorus, organic phosphate phosphorus, and total phosphate phosphorus.Inorganic nitrogen concentrations observed in this study were as high or higher than that previously reported for other bodies of water. Nitrate nitrogen entered the study area in relatively high concentrations from Comal Springs which was a major source of water for the Guadalupe River. Water from Canyon Reservoir, the other major source of water, was relatively low in nitrate nitrogen. The concentration of nitrate nitrogen was, therefore, dependent in part upon the portion of the total river flow originating from the two sources. Increased discharge from Canyon Reservoir and utilization by plants in areas of high chlorophyll a resulted in low nitrate-nitrogen levels. Retention of water in reservoirs reduced the concentration of nitrate nitrogen due to increased utilization by plants in areas of low flow. Nitrate nitrogen, in general, reached seasonal minima in summer and maxima in winter. Nitrite nitrogen showed considerable variation with no meaningful pattern except that higher concentrations occurred in association with high chlorophyll a and high Kjeldahl nitrogen, regions and periods of low river flow, and large phytoplankton populations. There was no increase in concentration of any form of nitrogen in the vicinity of sewage outfalls and no downstream accrual.Phosphorus levels in the study area were as high or higher than those reported in studies of other bodies of water. Sewage treatment plants at New Braunfels and Seguin, Texas, were major sources of phosphorus to the Guadelupe River. Total phosphate phosphorus was determined to be the most critical phosphate parameter in assessing eutrophication. Seasonally, it ranged from a winter high to a summer low. Concentrations were highest immediately below sewage outfalls and decreased as water progressed downstream. Inorganic-phosphate-phosphorus concentrations showed no clear seasonal trend but were clearly associated with sewage outfalls. Since large standing crops of phytoplankton were observed in areas of low inorganic phosphate phosphorus, it was not considered to limit photosynthesis. Total organic phosphate phosphorus varied seasonally, with high concentrations occurring during the spring and low concentrations in the fall. Total organic phosphate phosphorus showed no correlation with sewage outfalls, but was correlated to a degree with total Kjeldahl nitrogen and chlorophyll a. No consistent pattern of diel fluctuations was evident for any phosphorus or nitrogen compounds analyzed.     

Reduced Phosphorus Requirement of a Mutant Azolla-Anabaena Symbiotic N2-fixing Complex

The use of the photo-autotrophic nitrogen-fixing water fernAzolla as an effective source of organic nitrogen in tropicalpaddy fields has been limited by a high phosphorus requirement.Azolla species with a minimum of 1.5 to 2.0 mM phosphate (P)requirement, under controlled conditions, are known. A local Azolla species requiring at least 1.5 mM sodium phosphatefor a normal rate of multiplication and N₂ fixation was exposedto N-methyl-N'-nitro-N-nitrosoguanidine (MNNG). The resultingmutant population had a significantly lower P requirement, butwas auxotrophic for glutamine with an extremely reduced glutaminesynthetase (GS) activity. An L-methionine-DL-sulphoximine (MSX)-resistant(MSX^r) Azolla population, having an approximately 1.5 timeshigher GS activity than that of the wild type (WT) parent organism,was cultured and subjected to MNNG-induced mutation for lowP requirement while putting MSX as a control in the mutant selectionmedium. The resulting population of mutant Azolla was a normalprototroph with a P requirement as low as 0.75 mM for its ‘WTparent-like’ usual growth and N₂ fixation. Key words: Azolla, phosphorus requirement, mutation     

Does the chlorophyll a content of phytoplankton vary with trophic status in lakes on the New Zealand central volcanic plateau?

Recently, it has been shown that ratios of chlorophyll a toparticulate phosphorus (Chl a/PP) and chlorophyll a to particulatenitrogen (Chl a/PN) were significantly higher in eutrophic thanoligo/mesotrophic waters in 17 lakes on the central volcanicplateau, North Island, New Zealand. This difference was thoughtto be due to an increase in the chlorophyll a content of phytoplanktonin these eutrophic lakes. Corresponding measurements of chlorophylla and phytoplankton cell volume made during this study do notsupport this hypothesis. However, ratios of chlorophyll a toadenosine triphosphate and estimates of percentage phytoplanktonbiomass were significantly higher (P<0.05) in our eutrophicthan oligo/mesotrophic samples, suggesting that Chl a/PP andChl a/PN may be high in eutrophic waters simply because phytoplanktoncomprise more of the total microbial biomass. This hypothesisis supported by a strong linear relationship (r=0.88, P<0.001)between Chl a/PP and percentage phytoplankton biomass in sixof our study lakes where corresponding measurements were made.     

Subsurface phytoplankton blooms fuel pelagic production in the North Sea

The seasonal phytoplankton biomass distribution pattern in stratifiedtemperate marine waters is traditionally depicted as consistingof spring and autumn blooms. The energy source supporting pelagicsummer production is believed to be the spring bloom. However,the spring bloom disappears relatively quickly from the watercolumn and a large proportion of the material sedimenting tothe bottom following the spring bloom is often comprised ofintact phytoplankton cells. Thus, it is easy to argue that thespring bloom is fueling the energy demands of the benthos, butmore difficult to argue convincingly that energy fixed duringthe spring bloom is fueling the pelagic production occurringduring summer months. We argue here that periodic phytoplanktonblooms are occurring during the summer in the North Sea at depthsof >25 m and that the accumulated new production [sensu (Dugdaleand Goering, Limnol. Oceanogr., 12, 196–206, 1967)] occurringin these blooms may be greater than that occurring in the springbloom in the same regions. Thus, such blooms may explain apparentdiscrepancies in production yields between different temperatemarine systems.     

Phosphorus uptake by resting cysts of the marine dinoflagellate Scrippsiella trochoidea

Resting cysts of the marine dinoflagellate Scrippsiella trochoideawere produced under phosphorus (P)-deficient conditions, separatedfrom vegatative cells, and incubated for 28 days in darknessat 4 and 20°C in P-enriched and P-deplete medium. The Pcontent of cysts incubated in the P-replete medium was significantlyhigher than that of cysts in P-deplete medium. As the P contentof the cysts increased through time, dissolved inorganic phosphatewas depleted in the medium. This decrease cannot be attributedto free-living bacterial uptake, since there was no correspondingincrease in bacterial particulate P. Disappearance of P fromthe medium can, therefore, only be explained by uptake associatedwith the cysts. This could be either direct cyst uptake, uptakeby bacteria closely associated with the cysts, or adsorptionof P on the cyst wall. Evidence is strongest that the cystsincorporated phosphate during the resting stages of dormancyand quiescence, despite the fact that these are periods of significantlyreduced metabolism. Accumulation of P during these benthic restingstages would increase the survival of newly excysted vegetativecells as they re-enter the water column after germination, providinga competitive advantage over other phytoplankton. Freshwaterand marine sediments provide a P-rich environment which mayserve as a potential nutrient pool for dinoflagellate restingcysts. Mobilization of nutrients to and from the sediments viacysts must now be evaluated to ascertain whether this couldbe a significant term in nutrient budgets.     

Species-specific differences in phytoplankton responses to N and P enrichments and the N:P ratio in the Archipelago Sea, northern Baltic Sea

A nutrient enrichment experiment was conducted in order to studythe role of nitrogen (N), phosphorus (P) and the N:P ratio onthe early summer phytoplankton community in the ArchipelagoSea, northern Baltic Sea. The phytoplankton community was, interms of chlorophyll a and total biomass, primarily N-limited,but the individual species varied in their responses to thenutrient supply. The recorded overall N limitation was due tofast growth responses of a few N-limited species such as thediatom Chaetoceros wighamii (Brightwell) and the mixotrophicchrysophyte Uroglena sp. Another dominating diatom, Skeletonemacostatum (Greville) Cleve was most clearly P-limited. The N:Pratio had the strongest effect on Uroglena sp., which grew exponentiallyin the enrichments with a high N:P ratio. This can be explainedby the ability of the species to feed on P-rich bacteria, whichgives it a competitive advantage in P-limited conditions. Thespecies-specific differences in the responses to the nutrientenrichments can generally be explained by differences in thespecies physiology and they were consistent with the theoryof resource competition.