Size‐spectrum based differential response of phytoplankton to nutrient and iron‐organic matter combinations in microcosm experiments in a Chilean Patagonian Fjord

The Patagonian fjords have been recognized as a major region of relatively high primary productivity systems during spring–summer bloom periods, where iron‐organic matter forms may be essential complexes involved in key growth processes connected to the carbon and nitrogen cycles. We used two dissolved organic matter (DOM) types, marine polysaccharide and siderophore, as a model to understand how they affect the bioavailability of Fe to phytoplankton and bacteria and to assess their ecological role in fjord systems. A 10‐day microcosm study was performed in the Comau Fjord during summer conditions (March 2012). Pico‐, nano‐, and microphytoplankton abundance, total chlorophyll‐a and bacteria abundance, and bacterial secondary production estimates were analyzed in five treatments: (i) control (no additions), (ii) only nutrients (NUT: PO₄, NO₃, Si), (iii) nutrients + Fe(II), (iv) polysaccharide (natural diatoms extracted: 1–3 beta Glucan), and (v) Hexandentate Desferroxiamine B (DFB, siderophore). Our results showed that while DFB reduced Fe bioavailability for almost all phytoplankton assemblages in the fjord, polysaccharide did not have effects on the iron bioavailability. At Nutrients + Fe and Polysaccharide treatments, chlorophyll‐a concentration abruptly increased from 0.9 to 20 mg m^?3 during the first 4–6 days of the experimental period. Remarkably, at the Nutrients + Fe treatment, the development of the bloom was accompanied by markedly high abundances of Synechococcus, picoeukaryotes, and autotrophic nanoflagellates within the first 4 days of the experiment. Our study indicated that small plankton (phytoplankton <20 μm and bacteria) were the first to respond to dissolved Nutrients + Fe compared to large sized micro‐phytoplankton cells (>20 μm). This could be at least partially attributed to biological utilization of Fe (2 to 3 nM) by <20 μm phytoplankton and bacteria through the interaction with organic ligands released by bacteria that eventually could increase solubility of the Fe dissolved fraction thus having a positive effect on the small‐sized phytoplankton community.     

Grazing impact of microzooplankton on a diatom bloom in a mesocosm as estimated by pigment-specific dilution technique

To investigate the impact of microzooplankton grazing on phytoplankton bloom in coastal waters, an enclosure experiment was conducted in Saanich Inlet, Canada during the summer of 1996. Daily changes in the microzooplankton grazing rate on each phytoplankton group were investigated with the growth rates of each phytoplankton group from the beginning toward the end of bloom using the dilution technique with high-performance liquid chromatography (HPLC). On Day 1 when nitrate and iron were artificially added, chlorophyll a concentration was relatively low (4.3 μg l⁻¹) and 19′-hexanoyloxyfucoxanthin-containing prymnesiophytes were predominant in the chlorophyll biomass. However, both the synthetic rates and concentrations of 19′-hexanoyloxyfucoxanthin declined before bloom, suggesting that 19′-hexanoyloxyfucoxanthin-containing prymnesiophytes weakened. Chlorophyll a concentration peaked at 23 μg l⁻¹ on Day 4 and the bloom consisted of the small chain-forming diatoms Chaetoceros spp. (4 μm in cell diameter). Diatoms were secondary constituents in the chlorophyll biomass at the beginning of the experiment, and the growth rates of diatoms (fucoxanthin) were consistently high (>0.5 d⁻¹) until Day 3. Microzooplankton grazing rates on each phytoplankton group remarkably increased except on alloxanthin-containing cryptophytes after the nutrient enrichments, and peaked with >0.6 d⁻¹ on Day 3, indicating that >45% of the standing stock of each phytoplankton group was removed per day. Both the growth and mortality rates of alloxanthin-containing cryptophytes were relatively high (>1 and >0.5 d⁻¹, respectively) until the bloom, suggesting that a homeostatic mechanism might exist between predators and their prey. Overall, microzooplankton grazing showed a rapid response to the increase in phytoplankton abundance after the nutrient enrichments, and affected the magnitude of the bloom significantly. High grazing activity of microzooplankton contributed to an increase in the abundance of heterotrophic dinoflagellates with 7-24 μm in cell size, the fraction of large-sized (>10 μm) chlorophyll a, and stimulated the growth of larger-sized ciliates after the bloom.     

Food acquisition responses of the suspension-feeding bivalve Placopecten magellanicus to the flocculation and settlement of a phytoplankton bloom

In situ technologies were employed to monitor suspended particle flocculation and floc settlement and utilization by a cohort of sea scallops (Placopecten magellanicus) during the 2000 spring phytoplankton bloom in Bedford Basin, Nova Scotia, Canada. The objectives were to determine the effect of bloom flocculation and settling on food acquisition and utilization by scallops, and to assess the potential role of flocculation in enhancing the bioavailability of trophic resources and particle-reactive contaminants to bivalve filter feeders. The development and flocculation of the phytoplankton bloom were monitored within the surface layer (10 m depth) by in vivo chlorophyll fluorescence and silhouette camera observations. Sedimentation rate, seston abundance and composition, and sea scallop functional responses were monitored at 20 m depth (below the bloom) to provide insight into the potential forcing of feeding and digestion processes by changes in the abundance, composition and properties of the ambient food supply. The bloom began in mid-March and median floc diameter at 10 m depth increased rapidly from 200 μm to greater than 400 μm between 21 and 28 March. Flocs were observed to be abundant in the surface layer up to 4 April. Daily vertical particle flux was high during the last week of March and declined to near zero by 1 April. Clearance rates of scallops held at 20 m depth were relatively high (average ± S.D.; 11.7 ± 4.0 L h^− 1) during the period of bloom settlement and declined rapidly to low levels (0.4 ± 0.9 L h^− 1) after 31 March. Average absorption efficiency also declined (0.88 ± 0.01 to 0.78 ± 0.05) after bloom settlement. Daily biodeposition rates by scallops were poorly correlated with temporal variations in the quantity (total particulate matter and chlorophyll a concentration) or quality (organic content) of seston available to the scallops, but were significantly correlated with sedimentation rate. Comparison of disaggregated inorganic particle size distributions for suspended particulate matter, settled particles, and scallop feces indicated that fine-grained particles (1 to 4 μm) were effectively ingested by sea scallops—an indication of whole floc ingestion. The settlement of flocs produced during the spring bloom appears to be important in regulating this species physiological energetics and for enhancing the bioavailablility of fine particles (including picoplankton) and particle-reactive contaminants.     

Seasonal variation in abundance and size structure of phytoplankton in Baie des Chaleurs,southwestern Gulf of St. Lawrence,in relation to physical oceanographic conditions

We investigated changes in the abundance and size structure of phytoplankton and organic seston in relation to temperature, stratification and current patterns at Gascons on the north shore of Baie des Chaleurs, eastern Canada. Phytoplankton biomass showed a general decrease during the study (May to November 1989), except for a brief diatom bloom in late October. During most of the summer, a strong temperature driven stratification was present and <5 µm cells dominated the phytoplankton community. Particles measuring <5 µm also dominated the particulate organic matter (POC and PON) throughout the year. However, only 40% of these particles could be associated with phytoplankton cells. For both particulate matter and phytoplankton, the abundance of the <5 µm size fraction was positively correlated with the Brunt-Väilsälä index of stability of the water column. Inorganic nitrogen may have limited the phytoplankton growth, as generally reported for stratified environments. Most of the biomass was probably supported by nitrogen regenerated through microbial organisms. A large bacterioplankton community was suggested by the abundance of small (<5 µm) non-phytoplanktonic particles with a low and relatively uniform C/N ratio. Larger particles were only abundant at the beginning of the study (May–June) and on one date in October. Their C/N ratios indicated they were of varied origins.     

The role of phytoplankton in pollutant transfer processes in rivers. Example of River Marne (France)

The impact of the spring phytoplanktonic blooms on the partition of nutrients and metals between the dissolved and particulate phases was studied over two years in the river Marne, upstream of its confluence with the river Seine. Particular attention was devoted to copper and manganese, determined in the dissolved phase (< 0.22 m), subdivided into the colloidal phase and the 'truly dissolved' phase (< 10 KDaltons) obtained by tangential ultrafiltration. During the algal growth peaks, a sharp reduction of nutrients was observed in the dissolved phase: in 1994, consumption of 70% of silica, 50% of ammonium and 40% of soluble phosphorus present before the phytoplanktonic bloom.The water concentration peaks of suspended matter (SM) and of particulate metals, phosphorus and carbon coincide with those of chlorophyll. The highly significant correlations between pH and chlorophyll (r = 0.92, P < 0.001), pH and SM (r = 0.97, P < 0.001) and chlorophyll and SM (r = 0.93, P < 0.001) confirmed the phytoplanktonic nature of SM and the determining role of pH in the partition of metals, its increase being responsible for the coprecipitation of metal. As a corollary to the increase in the particulate phase, a decrease of 55% was observed for copper and manganese in the 'truly dissolved' phase compared to the concentrations outside bloom periods.The importance of the colloidal fraction was also shown. During the strong algal growth periods, the proportion of 'colloidal metal' in the dissolved phase (< 0.22 m) reached 60% and the increase in the colloidal fraction was 4 times stronger for copper than for manganese. While manganese seemed to be more associated with macroparticles, copper was more associated with fine colloidal biological particles intrinsic to the phytoplankton.     

Dynamics in the size structure of Skeletonema costatum (Greville) Cleve under conditions of reduced photosynthetically available radiation in a dredged tropical estuary

This study investigates the size-fractionated productivity and chlorophyll a concentrations in Ponggol estuary, a heavily dredged, light-limited and eutrophic tropical estuary located on the northeastern coast of Singapore. A 90% reduction in the photosynthetically available radiation (PAR) was seen in the subsurface waters of the dredged stations, when compared to an average reduction of about 75% in the subsurface waters of the un-dredged station. High phytoplankton production rates and chlorophyll a concentrations were recorded in the surface waters, with a significant reduction in the subsurface waters, especially at the two dredged stations. Out of the four size-classes of phytoplankton (0.2-2 μm, 2-20 μm, 20-200 μm and >200 μm) investigated, the relative dominance of the smaller size class of 2-20 μm over the 20-200 μm size class shifted depending on the availability of PAR. The size class 2-20 μm was observed to contribute up to 60 and 57% of the total production, respectively, in the surface waters of the un-dredged and dredged stations of the estuary. A relatively major contribution of 49% came from the cells of the small size-fraction of 2-20 μm in the subsurface waters of the un-dredged station. On the contrary, cells of the 20-200 μm size-class contributed up to 58% in the subsurface waters of the dredged station. Microscopic examination of the phytoplankton cells sampled showed morphological differences in the cells of the phytoplankton species Skeletonema costatum with some cells being distinctly larger in size than others. The larger cells were predominant in the dredged subsurface waters. In situ mesocosms with treatments exposed to high light irradiation registered a significant contribution by the smaller size-class of 2-20 μm, in contrast to the larger 20-200 μm dominating in the two treatments subjected to low light conditions. Based on the observations in this study and a review of the literature, it is hypothesized that the cells of S. costatum may actually be of two genetically different strains, whose relative dominance in the environment may be controlled by the quantum of available light. Thus, this shift in the relative dominance of one size fraction over the other is a response to altered PAR levels as the result of dredging.     

Colloidal organic carbon and trace metal (Cd, Fe, and Zn) releases by diatom exudation and copepod grazing

Colloidal macromolecular organic compounds are important intermediaries between solution and particle phases and play a critical role in the biogeochemistry of trace metals and organic carbon. The releases of colloidal organic carbon and trace metals (Cd, Fe, and Zn) mediated by copepod grazing and decomposition, and direct diatom exudation, were examined using a radiotracer approach. The colloidal phase was operationally defined in this study as the size fraction between 5 kDa and 0.2 μm and the dissolved phase as the ≤0.2 μm filter passing phase. About 13-60% of dissolved carbon exuded by the diatom Thalassiosira pseudonana was partitioned into the colloidal phase, and this fraction increased considerably as the diatom cells grew older. A lower fraction of dissolved ¹⁴C (12-23%) excreted by the copepods Acartia erythraea was detected in the colloidal phase compared to carcass (13-35%) and feces decomposition (21-34%). In contrast to carbon, a lower fraction of regenerated dissolved Cd (1-11%) and Zn (0-20%) from copepods and diatoms was consistently detected in the colloidal phases. Copepod excretion and carcass decomposition resulted in more colloidal Fe (51-91%) than diatom exudation (46-62% for Thalassiosira weissflogii, and 3-33% for T. pseudonana) and copepod feces decomposition (16-30%). Copepod (Calanus sinicus) grazing reduced the colloidal fraction of dissolved ¹⁴C, although a higher concentration of the diatom's (T. weissflogii) carbon was regenerated into the dissolved phase. The grazing of these copepods did not have any influence on the colloidal metal partitioning. The release of trace metals and carbon was enhanced by a higher density of copepod's grazing. Thus, different biological processes (grazing, excretion, exudation, and decomposition) may contribute differently to the production and dynamics of colloidal carbon and metals in planktonic systems.     

Effects of N and P supply on phytoplankton in Bizerte Lagoon (western Mediterranean)

Enriched bottle experiments were conducted in situ during winter (January and February) and summer (July and August) 2001 to examine the effects of nutrient enrichments (+ N, + P and + NP) on phytoplankton in Bizerte Lagoon, Tunisia. Chlorophyll a (Chl a), ranging from 3.05 μg L⁻¹ in winter to 4.52 μg L⁻¹ in summer, was dominated by the small size-faction (<5 μm) during both seasons. However, the contribution of the large size-fraction (5-200 μm) to Chl a increased from winter (26%) to summer (37%). Similarly, the carbon biomass of the 5-200 μm algae increased during the July/August period that was characterised by the high proliferation of several diatom taxa. In winter, N was the limiting element for phytoplankton growth. Its addition alone (+ N) or with P (+ NP) increased both the <5 μm and 5-200 μm Chl a concentrations. There was no change in the phytoplankton size structure, with the small cells dominating the final algal biomass in all treatments after 5 days. In summer, N and P limited the phytoplankton, but small and large algae exhibited diverse responses to different nutrient enrichments: addition of P increased the Chl a only in the 5-200 μm fraction, the + N treatment enhanced both size classes, and the NP fertilisation mostly stimulated the biomass of large cells. Consequently, the N and P addition in summer was followed by a significant change in the phytoplankton size structure, since both size-fractions contributed equally to the final Chl a biomass. Within the 5-200 μm algal community, various taxa had diverse responses to the nutrient supply during both seasons, leading to a change in the final community composition. The autotrophic flagellates appeared to grow well under N-deficient conditions. In contrast, diatom growth and biomass were mostly stimulated by the N enrichment while dinoflagellates exhibited the highest increase in their growth and biomass with P fertilisation. Our results suggest that the increasing anthropogenic supply of nutrients in the lagoon may influence algal dynamics as well as productivity in different ways depending on the nutrient composition.     

Biogeochemistry and effects of copper,manganese and zinc added to enclosures in Island Billabong,Magela Creek,northern Australia

Three large plastic enclosures (5 m diam, volume 40 m³) were used to study the effects of copper, manganese and zinc, on the phytoplankton community in Island billabong, a floodplain billabong (waterhole) situated in the Magela Creek in tropical northern Australia. Copper was added to one enclosure, and manganese and zinc to another, to give initial concentrations around ten times the normal wet season values. The enclosures and the billabong were monitored over a ten week period towards the end of the dry season, with the enclosures allowed to stabilise for four weeks before the metals were added.The control enclosure adequately simulated the temperature and pH changes in the billabong. The trends in conductivity, dissolved oxygen and major ion concentrations were similar in the enclosure and the billabong, with the minor differences observed attributed to either epiphytic growth on the enclosure walls (influenced dissolved oxygen, pH and bicarbonate concentration) or ingress of sulphate-rich groundwater into the billabong (influenced sulphate concentration and conductivity). Major differences in both the composition of species and the size of the phytoplankton populations were observed between the three enclosures and between the control enclosure and the billabong. This variability reflects the great natural variability in the phytoplankton communities in tropical lentic systems, and means that enclosures are unlikely to adequately simulate the biological communities in the billabongs.The control enclosure appeared to simulate quite well the longer term changes in total concentration and speciation of the three metals (copper, manganese & zinc) in the billabong. The mean concentrations of copper and zinc were similar in the two systems, although the mean concentration of manganese in the billabong was almost double that in the enclosure, possibly due to ingress of manganese-enriched groundwater. Particulate forms dominated the speciation of copper and manganese. There was considerable short term variation in both total metal concentration and speciation in both the enclosure and the billabong. This variability appears to be a feature of these small tropical waterbodies.The added heavy metals were found to have minimal detrimental effect on the phytoplankton community in each metal-loaded enclosure. The high natural variability in the phytoplankton community in these tropical systems will make it difficult to separate natural changes from those caused by low level contamination from mining operations should this occur.All three metals were rapidly removed from the water column, so that by the end of the six week period, only ca. 5% of each added metal remained in the water column. Association with the particulate matter (phytoplankton, abiotic particulate matter and MnO_x in enclosure 2) followed by sedimentation was the major removal pathway. Epiphytes growing on the enclosure walls appeared to have a minor influence (<10% of the total amount of metal added) on the removal of the added metals. For copper, uptake by phytoplankton followed by sedimentation was the major (65%) removal process. Manganese and zinc, added together, were found to influence each other. The major manganese removal process (60%) was rapid (ca. 3 days) involving bacterial oxidation and sedimentation of the MnO_x formed. This material appeared to have little influence on the behaviour of zinc, possibly because other particulate matter competed more effectively for the zinc. A further 30% of the added manganese was removed via initial adsorption to other particulate matter, possibly phytoplankton. Approximately one third of this adsorbed manganese (10% of the total added) appeared to undergo delayed oxidation some 8 days after the initial additions, and the heavier particles settled out more rapidly. This path was responsible for removing the major amount (ca. 60%) of the added zinc. We hypothesis that the sorbed zinc inhibited the bacterial oxidation of the manganese. A further 25% of the zinc was removed in association with a burst of phytoplankton activity. The occurrence of bursts in the phytoplankton activity, when populations can increase very substantially and then decrease again, all within the space of a day, appears to be an important mechanism for removing copper and zinc from the water column in these tropical water bodies     

Summer diatom blooms in the North Pacific subtropical gyre: 2008-2009

The summertime North Pacific subtropical gyre has widespread phytoplankton blooms between Hawaii and the subtropical front (～30°N) that appear as chlorophyll (chl) increases in satellite ocean color data. Nitrogen-fixing diatom symbioses (diatom-diazotroph associations: DDAs) often increase 10(2)-10(3) fold in these blooms and contribute to elevated export flux. In 2008 and 2009, two cruises targeted satellite chlorophyll blooms to examine DDA species abundance, chlorophyll concentration, biogenic silica concentration, and hydrography. Generalized observations that DDA blooms occur when the mixed layer depth is < 70 m are supported, but there is no consistent relationship between mixed layer depth, bloom intensity, or composition; regional blooms between 22-34°N occur within a broader temperature range (21-26°C) than previously reported. In both years, the Hemiaulus-Richelia and Rhizosolenia-Richelia DDAs increased 10(2)-10(3) over background concentrations within satellite-defined bloom features. The two years share a common trend of Hemiaulus dominance of the DDAs and substantial increases in the >10 μm chl a fraction (～40-90+% of total chl a). Integrated diatom abundance varied 10-fold over <10 km. Biogenic silica concentration tracked diatom abundance, was dominated by the >10 μm size fraction, and increased up to 5-fold in the blooms. The two years differed in the magnitude of the surface chl a increase (2009>2008), the abundance of pennate diatoms within the bloom (2009>2008), and the substantially greater mixed layer depth in 2009. Only the 2009 bloom had sufficient chl a in the >10 μm fraction to produce the observed ocean color chl increase. Blooms had high spatial variability; ocean color images likely average over numerous small events over time and space scales that exceed the individual event scale. Summertime DDA export flux noted at the Hawaii time-series Sta. ALOHA is probably a generalized feature of the eastern N. Pacific north to the subtropical front.     

Seasonal changes in particulate and dissolved lipids in a eutrophic prairie lake

1. Critical periods of lipid energy transfer from phyto- to zooplankton were inferred by comparing seasonal patterns of particulate and dissolved lipid fractions in lake water with temporal changes in lipid energy reserves of the zooplankton in a hypereutrophic lake.
2. The midsummer phytoplankton community was dominated by the bloom-forming cyanobacterium Aphanizomenon flos-aquae. The collapse of the bloom was accompanied by a 2-week period of severe nitrogen deficiency after which there was a marked increase in the concentration of lipid energy reserves in the particulate (algal) fraction.
3. Areal lipid energy reserves of the dominant herbivorous zooplankton responded positively to changes in the tri-and diacylglycerol content of the particulate fraction of lake water in a species-specific manner.
4. Bacterial numbers also peaked in September concomitant with a large increase in free fatty acids in the dissolved lipid fraction probably produced by the decay of the
A. flos-aquae bloom.
5. The association between periods of nitrogen deficiency and increased energy reserve lipids in the particulate fraction supports observations made with laboratory algal cultures that periods of nutrient deficiency may intensify lipid synthesis in some algal species, thereby enhancing the rate of lipid energy transfer from phytoplankton to zooplankton.     

Size-dependent changes in phytoplankton C and N uptake in the dynamic mixed layer of Lake Biwa

1. Short-term (days) hydrodynamic effects of wind-induced mixing on phytoplankton size structure, and C and N uptake characteristics, were studied in the surface mixed layer (epilimnion) of Lake Biwa (North Basin), before and during a period of high winds (typhoons). 2. The latter period was characterized by two major typhoon events associated with deepening of the seasonal thermocline, reduced water column stability, decreased underwater irradiance and increased dissolved reactive N and particulate P. 3. Nutrient concentrations, seston C/N ratios, and uptake rates indicated that phytoplankton biomass and production were limited by P and not N throughout the study. Higher C- and N-based productivity during the typhoon period than before reflected the increased phytoplankton biomass and higher specific uptake rates due to increased nutrient supply. 4. Changes in the size-structure of phytoplankton (< 2 and > 2 μm) were associated with variations in the stratification and mixing regime. When vertical stability was high (before the typhoons) concentrations of > 2 μm biomass (chlorophyll a, particulate organic C and N) were higher at the bottom of the mixed layer than at the surface whereas, when stability of the mixed layer was low (the typhoon period), the contribution of picoplankton (< 2 μm) to total Chl a increased at the surface and decreased at the bottom following the first high winds. 5. Photoadaptive adjustments of the phytoplankton provided further evidence of hydrodynamic control. The lower intracellular Chl a concentrations and C and N uptake efficiencies in the < 2 μm fraction suggest that they experienced, on average, higher irradiance than the larger cells because of their lower sinking rates. During the stability period, picoplankton exhibited higher photosynthetic efficiencies at the bottom of the mixed layer than at the surface. Such differences disappeared during the typhoon period indicating that the mixing rate was then probably higher than the photoacclimation rate in the small size fraction. 6. The present results stress the highly transient nature of biological homogeneity in the surface mixed layer of the lake.     

Nitrate uptake by size-fractionated phytoplankton on the Scotian Shelf (Northwest Atlantic): spatial and temporal variability

New (nitrate) phytoplankton production was estimated monthlyduring 1 year (March 1991–March 1992) at three stationson the Scotian Shelf, Northwest Atlantic. Samples were sizefractionated to assess the uptake of nitrate by small (<5µm) and large (>5 µm) phytoplankton. The biomassof small phytoplankton remained relatively constant over theyear, whereas that of the large size fraction was high in earlyspring and low during the remainder of the year. Monthly variationsin nitrate uptake were similar for the two size fractions, suggestingthat both small and large phytoplankton used nitrate when available.It follows that, outside the spring bloom, new production waslargely due to the small fraction. Our results do not supportthe notion that new production is associated with large phytoplanktonand regenerated production with small phytoplankton.     

Temporal variations in phytoplankton, particulates, and colored dissolved organic material based on optical properties during a Long Island brown tide compared to an adjacent embayment

Since 1985, the coastal embayments of Long Island, New York, have been plagued with recurrent blooms, aptly called brown tides, of the pelagophyte Aureococcus anophagefferens. The distinct ocean color observed during these blooms suggests that optical methods can be used as a tool to study, detect, and track brown tides. Thus, the goal of our project was to compare the optical properties and pigment composition during bloom and non-bloom conditions and assess temporal variations in the phytoplankton and other constituents in the seawater associated with bloom development. From 17 May to 8 June 2000, we measured a time series of particle size distributions and concentrations as well as size-fractioned algal pigments and optical properties in two Long Island embayments where brown tides are known to occur. During our study, A. anophagefferens represented an insignificant contribution to the algal community in West Neck Bay (WNB), whereas a bloom developed in Quantuck Bay (QB). Initially, temperature and salinity were similar at the two locations; however, bulk optical properties, chlorophyll, and particle concentrations were nearly a factor of 2 greater at QB. Bulk optical properties remained constant at WNB, yet increased exponentially at QB as the bloom developed. The composition of particulates, including phytoplankton, varied little at QB, and the optical properties suggested the dominance of A. anophagefferens (confirmed by microscopy). The largest temporal variations were observed in the colored dissolved organic material (CDOM); the colloidal (0.2–0.7 μm) fraction, exhibiting a strong protein-like signal, increased dramatically at the height of the bloom. At WNB particle sizes and algal composition varied despite the invariant bulk optical properties; CDOM variations were minimal. Overall, the optical properties in the two bays demonstrated that at QB temporal variations were dominated by biomass and colloidal protein changes, whereas shifts in the algal community occurred at WNB. This study demonstrates the utility of in situ optical observations to resolve temporal changes in the ecological conditions associated with algal bloom development.     

Seasonal variations in upwelling and in the grazing impact of copepods on phytoplankton off A Coruña (Galicia, NW Spain)

The impact of grazing by copepods on phytoplankton was studied during a seasonal cycle on the Galician shelf off A Coruña (NW Spain). Grazing was estimated by measuring the chlorophyll gut content and the evacuation rates of copepods from three mesh-size classes: 200-500 (small), 500-1000 (medium), and 1000-2000 μm (large). Between February 1996 and June 1997, monthly measurements of water temperature, chlorophyll concentration, primary production rates, and copepod abundance, chlorophyll gut content, and evacuation rates were taken at an 80-m-deep, fixed shelf station. Additionally, the same measurements were collected daily during two bloom events in March and in July 1996. Small copepods were the most abundant through the seasonal cycle. The highest grazing impact, however, was due to the medium and large size classes. Grazing by small copepods exceeded grazing by medium and large copepods only during phytoplankton spring blooms. The impact of copepod grazing (considering all size fractions) was generally low. On average, 2% of the phytoplankton biomass and 6% of the primary production were removed daily by the copepod community. Maximum grazing impact values (9% of the phytoplankton biomass and 39% of the primary production) were found in mid-summer. These results suggest that most of the phytoplankton biomass would escape direct copepod grazing in this upwelling area.     

Size-fractionated phytoplankton chlorophyll in an Eastern Mediterranean coastal system (Maliakos Gulf, Greece)

The dynamics of phytoplankton biomass were studied in an Eastern Mediterranean semi-enclosed coastal system (Maliakos Gulf, Aegean Sea), over 1 year. In particular, chlorophyll a (chl a) was fractionated into four size classes: picoplankton (0.2–2 μm), nanoplankton (2–20 μm), microplankton (20–180 μm) and net phytoplankton (>180 μm). The spatial and temporal variation in dissolved inorganic nutrients and particulate organic carbon (POC) were also investigated. The water column was well mixed throughout the year, resulting in no differences between depths for all the measured parameters. Total chl a was highest in the inner part of the gulf and peaked in winter (2.65 μg l^–1). During the phytoplankton bloom, microplankton and net phytoplankton together dominated the autotrophic biomass (67.2–95.0% of total chl a), while in the warmer months the contribution of pico- and nanoplankton was the most significant (77.5–93.4% of total chl a). The small fractions, although showing low chl a concentrations, were important contributors to the POC pool, especially in the outer gulf. No statistically significant correlations were found between any chl a size fraction and inorganic nutrients. For most of the year, phytoplankton was not limited by inorganic nitrogen concentrations. Electronic Publication     

Changes in the plankton community following introduction of filter-feeding planktivorous fish

1. We conducted enclosure experiments in a shallow eutrophic lake, in which a biomass gradient of the filter-feeding planktivore, silver carp, Hypophthalmichthys molitrix Valenciennes, was created, and subsequent community changes in both zooplankton and phytoplankton were examined.
2. During a summer experiment, a bloom of Anabaena flos-aquae developed (≈ 8000 cells mL⁻¹) solely in an enclosure without silver carp. Concurrent with, or slightly preceding the Anabaena bloom, the number of rotifer species and their abundance increased from seven to twelve species (1700–14 400 organisms L⁻¹) after the bloom in this fish-free enclosure. Protozoans and bacteria were generally insensitive to the gradient of silver carp biomass.
3. During an autumn experiment, on the other hand, large herbivorous crustaceans were more efficient than silver carp in suppressing the algae, partly because the lower water temperature (≈ 24 °C) inhibited active feeding of this warm-water fish and also formation of algal colonies. Heterotrophic nanoflagellate and bacterial densities were also influenced negatively by the crustaceans.
4. Correspondence analysis (CA) was applied to the weekly community data of zooplankton and phytoplankton. A major effect detected in the zooplankton community was the presence/absence of silver carp rather than the biomass of silver carp, whereas that in the phytoplankton community was the fish biomass before the Anabaena bloom, but shifted to the presence/absence of the fish after the bloom.     

The submicron size fraction of seawater containing high numbers of virus particles as bioactive agent in unicellular plankton community successions

The effect of moderately (2.5 times) increasing the concentrationof the submicron size fraction (2–200 nm) of seawateron unicellular heterotrophic plankton organisms was tested ina series of experiments during algal bloom conditions. Usingelectron microscopy, vims-like particles (VLPs) were by farthe most visible component in this size fraction. We incubatedseawater. which was collected in late winter, in slowly rotatingcylindrical tanks that were exposed to spring temperature andlight conditions. This led to the onset of a mixed diatom bloomwhich was delayed by several days in submicron size-enrichedtreatments. Later, phytoplankton biomass reached values evenslightly higher than in unaltered incubations. The abundanceof heterotrophic unicellular plankton and VLPs exhibited morepronounced oscillation amplitudes during the early phase ofsuccession, thus being stimulated by additional submicron-sizedmaterial (including virions). VLPs are suggested as a possiblesupplementary diet for heterotrophic flagellates. From 190 honwards, toward the end of the incubation experiments (550 h).heterotrophic flagellate density was no longer influenced byinitial enrichment with the submicron size fraction. In contrast,bacteria were consistently repressed in the last two-thirdsof the experimental duration. Our results support the existenceof bioactive substances in the 2-200 nm size fraction, mostprobably virus particles, which are capable of influencing thedynamics and oscillation patterns in plankton community successionaccording to their concentration during algal bloom events.     

A new approach to estimate the in situ fractional degradation rate of organic matter and nitrogen in wheat yeast concentrates

In the classic in situ method, small particles are removed during rinsing and hence their fractional degradation rate cannot be determined. A new approach was developed to estimate the fractional degradation rate of nutrients in small particles. This approach was based on an alternative rinsing method to reduce the particulate matter loss during rinsing and on quantifying the particulate matter loss that occurs during incubation in the rumen itself. To quantify particulate matter loss during incubation, loss of small particles during the in situ incubation was studied using undegradable silica with different particle sizes. Particulate matter loss during incubation was limited to particles smaller than ~40 μm with a mean fractional particulate matter loss rate of 0.035 h⁻¹ (first experiment) and 0.073 h⁻¹ (second experiment) and an undegradable fraction of 0.001 and 0.050, respectively. In the second experiment, the fractional particulate matter loss rate after rinsing in a water bath at 50 strokes per minute (s.p.m.) (0.215 h⁻¹) and the undegradable fraction at 20 s.p.m. (0.461) were significantly larger than that upon incubation in the rumen, whereas the fractional particulate matter loss rate (0.140 and 0.087 h⁻¹, respectively) and the undegradable fraction (0.330 and 0.075, respectively) after rinsing at 30 and 40 s.p.m. did not differ with that upon rumen incubation. This new approach was applied to estimate the in situ fractional degradation rate of insoluble organic matter (OM) and insoluble nitrogen (N) in three different wheat yeast concentrates (WYC). These WYC were characterised by a high fraction of small particles and estimating their fractional degradation rate was not possible using the traditional washing machine rinsing method. The new rinsing method increased the mean non-washout fraction of OM and N in these products from 0.113 and 0.084 (washing machine method) to 0.670 and 0.782, respectively. The mean effective degradation (ED) without correction for particulate matter loss of OM and of N was 0.714 and 0.601, respectively, and significant differences were observed between the WYC products. Applying the correction for particulate matter loss reduced the mean ED of OM to 0.676 (30 s.p.m.) and 0.477 (40 s.p.m.), and reduced the mean ED of N to 0.475 (30 s.p.m.) and 0.328 (40 s.p.m.). These marked reductions in fractional degradation rate upon correction for small particulate matter loss emphasised the pronounced effect of correction for undegraded particulate matter loss on the fractional disappearance rates of OM and N in WYC products.     

Qualitative importance of the microbial loop and plankton community structure in a eutrophic lake during a bloom of cyanobacteria

Plankton community structure and major pools and fluxes of carbon were observed before and after culmination of a bloom of cyanobacteria in eutrophic Frederiksborg Slotssø, Denmark. Biomass changes of heterotrophic nanoflagellates, ciliates, microzooplankton (50 to 140 μm), and macrozooplankton (larger than 140 μm) were compared to phytoplankton and bacterial production as well as micro- and macrozooplankton ingestion rates of phytoplankton and bacteria. The carbon budget was used as a means to examine causal relationships in the plankton community. Phytoplankton biomass decreased and algae smaller than 20 μm replacedAphanizomenon after the culmination of cyanobacteria. Bacterial net production peaked shortly after the culmination of the bloom (510 μg C liter^?1 d^?1 and decreased thereafter to a level of approximately 124 μg C liter^?1 d^?1. Phytoplankton extracellular release of organic carbon accounted for only 4–9% of bacterial carbon demand. Cyclopoid copepods and small-sized cladocerans started to grow after the culmination, but food limitation probably controlled the biomass after the collapse of the bloom. Grazing of micro- and macrozooplankton were estimated from in situ experiments using labeled bacteria and algae. Macrozooplankton grazed 22% of bacterial net production during the bloom and 86% after the bloom, while microzooplankton (nauplii, rotifers and ciliates larger than 50 μm) ingested low amounts of bacteria and removed 10–16% of bacterial carbon. Both macro-and microzooplankton grazed algae smaller than 20 μm, although they did not control algal biomass. From calculated clearance rates it was found that heterotrophic nanoflagellates (40–440 ml^?1) grazed 3–4% of the bacterial production, while ciliates smaller than 50 μm removed 19–39% of bacterial production, supporting the idea that ciliates are an important link between bacteria and higher trophic levels. During and after the bloom ofAphanizomenon, major fluxes of carbon between bacteria, ciliates and crustaceans were observed, and heterotrophic nanoflagellates played a minor role in the pelagic food web.