The herbivorous impact of microzooplankton during two short-term Lagrangian experiments off the NW coast of Galicia in summer 1998

Microzooplankton (heterotrophic microplankton and heterotrophic nanoflagellates) and their herbivorous activity were estimated from dilution experiments in August 1998 during two Lagrangian drift experiments that sampled contrasting conditions—an upwelling/relaxation event along the shelf edge and an oligotrophic offshore filament. During upwelling/relaxation, heterotrophic microplankton were present at mean surface concentrations between 15,000 and 48,000 cells l⁻¹. Heterotrophic nanoflagellate concentrations were between 200 and 700 cells ml⁻¹ and the most abundant component of the heterotrophic microplankton was the aloricate choreotrich ciliates which increased dramatically in concentration from 6,000 to 24,000 cells l⁻¹ during the first 4 days of the study. Total microzooplankton biomass reached a maximum of 39mgC.m⁻³. In the filament, which developed from the upwelling, cell concentrations were lower and averaged 4,500 cells l⁻¹ for heterotrophic microplankton and 250 cells ml⁻¹ for heterotrophic nanoflagellates. Total microzooplankton biomass was about 10–12mgC.m⁻³. Microzooplankton turned over between 40 and 85% of the phytoplankton standing stock, thereby consuming between 5 and 78mg phytoplankton carbon.m⁻³.d⁻¹. The magnitude of this activity was highest during upwelling/relaxation and was positively correlated to heterotrophic nanoflagellate biomass and chlorophyll-a concentration but not heterotrophic microplankton biomass. The proportion of primary production grazed decreased from 160 to 59% d⁻¹ during upwelling/relaxation and ranged between 60 and 90% d⁻¹ in the filament. Microzooplankton herbivory within the euphotic zone increased from 684 to >2000mgC.m⁻².d⁻¹ during upwelling/relaxation and was between 327 and 802mgC.m⁻².d⁻¹ in the filament. Although microzooplankton herbivory was lower and less variable during the filament study, microzooplankton consumed on average 60% of the phytoplankton standing stocks which was higher than found during upwelling/relaxation. Microzooplankton assimilation efficiency ranged between 3 and 33% during upwelling/relaxation and between 0 and 13% in the filament. Our data demonstrate a close coupling between phytoplankton growth and microzooplankton herbivory in surface waters off the Galician Coast and suggest that microzooplankton may have been a significant sink for phytogenic carbon during August 1998.     

Phytoplankton growth, microzooplankton herbivory and community structure in the southeast Bering Sea: insight into the formation and temporal persistence of an Emiliania huxleyi bloom

Using the seawater dilution technique, we measured phytoplankton growth and microzooplankton grazing rates within and outside of the 1999 Bering Sea coccolithophorid bloom. We found that reduced microzooplankton grazing mortality is a key component in the formation and temporal persistence of the Emiliania huxleyi bloom that continues to proliferate in the southeast Bering Sea. Total chlorophyll a (Chl a) at the study sites ranged from 0.40 to 4.45 μg C l⁻¹. Highest phytoplankton biomass was found within the bloom, which was a mixed assemblage of diatoms and E. huxleyi. Here, 75% of the Chl a came from cells >10 μm and was attributed primarily to the high abundance of the diatom Nitzschia spp. Nutrient-enhanced total phytoplankton growth rates averaged 0.53 d⁻¹ across all experimental stations. Average growth rates for >10 μm and <10 μm cells were nearly equal, while microzooplankton grazing varied among stations and size fractions. Grazing on phytoplankton cells >10 μm ranged from 0.19 to 1.14 d⁻¹. Grazing on cells <10 μm ranged from 0.02 to 1.07 d⁻¹, and was significantly higher at non-bloom (avg. 0.71 d⁻¹) than at bloom (avg. 0.14 d⁻¹) stations. Averaged across all stations, grazing by microzooplankton accounted for 110% and 81% of phytoplankton growth for >10 and <10 μm cells, respectively. These findings contradict the paradigm that microzooplankton are constrained to diets of nanophytoplankton and strongly suggests that their grazing capability extends beyond boundaries assumed by size-based models. Dinoflagellates and oligotrich ciliates dominated the microzooplankton community. Estimates of abundance and biomass for microzooplankton >10 μm were higher than previously reported for the region, ranging from 22,000 to 227,430 cells l⁻¹ and 18 to 164 μg C l⁻¹. Highest abundance and biomass occurred in the bloom and corresponded with increased abundance of the large ciliate Laboea, and the heterotrophic dinoflagellates Protoperidinium and Gyrodinium spp. Despite low grazing rates on phytoplankton <10 μm within the bloom, the abundance and biomass of small microzooplankton (<20 μm) capable of grazing E. huxleyi was relatively high at bloom stations. This body of evidence, coupled with observed high grazing rates on large phytoplankton cells, suggests the phytoplankton community composition was strongly regulated by herbivorous activity of microzooplankton. Because grazing behavior deviated from size-based model predictions and was not proportional to microzooplankton biomass, alternate mechanisms that dictate levels of grazing activity were in effect in the southeastern Bering Sea. We hypothesize that these mechanisms included morphological or chemical signaling between phytoplankton and micrograzers, which led to selective grazing pressure.     

The contribution of microzooplankton to the diet of mesozooplankton in an upwelling filament off the north west coast of Spain

Incubation experiments were carried out daily during a Lagrangian experiment within an upwelled filament off the Galician coast to determine the importance of microzooplankton in the diet of calanoid copepods. Despite low chlorophyll concentrations the microzooplankton formed the minor component of the diet of the copepod community (7 to 15% of carbon ingested through autotrophic and heterotrophic prey). Ingestion of ciliates was greater than that of heterotrophic dinoflagellates, which reflected a higher abundance of ciliates in the water column. Heterotrophic nanoflagellates appeared also to be consumed by the copepods, although the very small size fraction (2–5μm) was probably not grazed by the larger copepods of Calanus spp. Grazing pressure by the copepods enumerated in the net samples was not sufficient to impact significantly the microzooplankton populations (2 to 51% of daily microzooplankton production was removed). Allometric relationships of grazing on microzooplankton for a range of numerically dominant copepod species are developed from the experimental results. The grazing pressure of the whole copepod community is estimated from these relationships. By considering the total mesozooplankton community we suggest that microzooplankton growth was probably restricted by metazoan grazers.     

Primary production and nutrient assimilation in the Iberian upwelling in August 1998

Primary production was measured during two Lagrangian experiments in the Iberian upwelling. The first experiment, in a body of upwelled water, measured day-to-day changes in phytoplankton activity as the water mass moved south along the shelf break. Nutrient concentrations decreased over a five day period, with concomitant increases in phytoplankton biomass. Initially the maximum phytoplankton biomass was in the upper 10m but after four days, a sub-surface chlorophyll maximum was present at 30m. Depth-integrated primary production at the beginning of the experiment was 70mmolC.m⁻².d⁻¹ (838mgC.m⁻².d⁻¹) and reached a maximum of 88mmolC.m⁻².d⁻¹ (1053mgC.m⁻².d⁻¹) on day 3. On day 1, the picoplankton fraction (<2μm) was slightly more productive than larger (>5μm) phytoplankton, but the increase in overall production during the drift experiment was by these larger cells. Nitrate was the dominant nitrogen source. As nutrient concentrations declined, ammonium became increasingly more important as a nitrogen source and the f-ratio decreased from 0.7 to 0.5. Picoplankton cells (<2μm) were responsible for most (65–80%) of the ammonium uptake. The C:N:P uptake ratios were very close to the Redfield ratio for the first four days but as nutrients became depleted high C:N uptake ratios (11 to 43) were measured. Over the period of the experiment, nitrate concentration within the upper 40m decreased by 47.91mmolN.m⁻². In vitro estimates, based on ¹⁵N nitrate uptake, accounted for 56% of the decrease in nitrate concentration observed in the drifting water mass. Ammonium uptake over the same four day period was 16.28mmolN.m⁻², giving a total nitrogen uptake of 43.18mmolN.m⁻².In the second experiment, an offshore filament was the focus and a water mass was sampled as it moved offshore. Nutrient concentrations were very low (nitrate was <10nmol l⁻¹ and ammonium was 20–40nmol l⁻¹). Primary production rate varied between 36mmolC.m⁻².d⁻¹ (436mgC.m⁻².d⁻¹) and 21mmolC.m⁻².d⁻¹ (249mgC.m⁻².d⁻¹). Picophytoplankton was the most productive fraction and was responsible for a constant proportion (ca 0.65) of the total carbon fixation. Uptake rates of both nitrate and ammonium were between 10 and 20% of those measured in the upwelling region. Urea could be a very significant nitrogen source in these waters with much higher uptake rates than nitrate or ammonium; urea turnover times were ca. one day but the source of the urea remains unknown. Urea uptake had a profound effect on calculated f ratios. If only nitrate and ammonium uptake was considered, f ratios were calculated to be 0.42–0.46 but inclusion of urea uptake reduced the f ratio to <0.1. The primary production of this oligotrophic off-shore filament was driven by regenerated nitrogen.     

Abundance, variability, and potential grazing impact of planktonic ciliates in the open subaratic Pacific Ocean

The abundance and variability of planktonic ciliates in the open subarctic Pacific were determined during four month-long cruises in 1987 and 1988. The ciliate community, numerically dominated by relatively small aloricate choreotrichs, was comparable in abundance to communities in a range of oceanic and neritic environments, including waters with much higher average chlorophyll concentrations. Integrated (0–80m) ciliate biomass was typically 100–200mgC m⁻², although 3- to 4-fold higher levels were observed on two occasions in spring. Ciliate community biomass, in general, was dominated by large (>20μm width) individuals, although in August 1988 the biomass of smaller cells was as great or greater. The estimated grazing impact of the ciliate community averaged 20% of the primary production. On one instance in May 1988, however, a large biomass of ciliates led to an estimated grazing impact equivalent to 55% of phytoplankton production. While ciliates may be major phytoplankton grazers during sporadic ciliate “blooms”, dino- and other heterotrophic flagellates, which make up the bulk of microheterotroph biomass, must normally be of equal or greater importance as herbivores in this ocean region.     

Seasonal variation in the community and size structure of nano- and microzooplankton in Gyeonggi Bay,Yellow Sea

To investigate the seasonal variation and community structure of nano- and microzooplankton in Gyeonggi Bay of the Yellow Sea, the abundance and carbon biomass of nano- and microzooplankton were evaluated at 10-day intervals from January 1997 to December 1999. Four major groups of nano- and microzooplankton communities were classified: heterotrophic ciliates, heterotrophic dinoflagellates (HDF), heterotrophic nanoflagellates (HNF), and copepod nauplii. The total carbon biomass of nano- and microzooplankton ranged from 10.2 to 168.8 μg C L⁻¹ and was highest during or after phytoplankton blooms. Nano- and microzooplankton communities were composed of heterotrophic ciliates (7.4–81.4%; average 41.7% of total biomass), HDF (0.1–70.3%; average 26.1% of total biomass), copepod nauplii (1.6–70.6%; average 20.7% of total biomass), and HNF (0.8–59.5%; average 11.5% of total biomass). The relative contribution of individual components in the nano- and microzooplankton communities appeared to differ by seasons. Ciliates accounted for the most major component of nano- and microzooplankton communities, except during summer and phytoplankton blooming seasons, whereas HDF were more dominant during the phytoplankton blooming seasons. The abundance and biomass of nano- and microzooplankton generally followed the seasonal dynamics of phytoplankton. The size and community distribution of nano- and microzooplankton was positively correlated with size-fractionated phytoplankton. The carbon requirement of microzooplankton ranged from 60 to 83% of daily primary production, and was relatively high when phytoplankton biomass was high. Therefore, our result suggests that the seasonal variation in the community and size composition of nano- and microzooplankton appears to be primarily governed by phytoplankton size and concentration as a food source, and their abundance may greatly affect trophic dynamics by controlling the seasonal abundance of phytoplankton.     

Grazing by meso- and microzooplankton on phytoplankton in the upper reaches of the Schelde estuary (Belgium/The Netherlands)

In contrast with the marine reaches of estuaries, few studies have dealt with zooplankton grazing on phytoplankton in the upper estuarine reaches, where freshwater zooplankton species tend to dominate the zooplankton community. In spring and early summer 2003, grazing by micro- and mesozooplankton on phytoplankton was investigated at three sites in the upper Schelde estuary. Grazing by mesozooplankton was evaluated by monitoring growth of phytoplankton in 200 μm filtered water in the presence or absence of mesozooplankton. In different experiments, the grazing impact was tested of the calanoïd copepod Eurytemora affinis, the cyclopoid copepods Acanthocyclops robustus and Cyclops vicinus and the cladocera Chydorus sphaericus, Moina affinis and Daphnia magna/pulex. No significant grazing impact of mesozooplankton in any experiment was found despite the fact that mesozooplankton densities used in the experiments (20 or 40 ind. l⁻¹) were higher than densities in the field (0.1–6.9 ind. l⁻¹). Grazing by microzooplankton was evaluated by comparing growth of phytoplankton in 30 and 200 μm filtered water. Microzooplankton in the 30–200 μm size range included mainly rotifers of the genera Brachionus, Trichocerca and Synchaeta, which were present from 191 to 1777 ind. l⁻¹. Microzooplankton had a significant grazing impact in five out of six experiments. They had a community grazing rate of 0.41–1.83 day⁻¹ and grazed up to 84% of initial phytoplankton standing stock per day. Rotifer clearance rates estimated from microzooplankton community grazing rates and rotifer abundances varied from 8.3 to 41.7 μl ind.⁻¹ h⁻¹. CHEMTAX analysis of accessory pigment data revealed a similar phytoplankton community composition after incubation with and without microzooplankton, indicating non-selective feeding by rotifers on phytoplankton.     

Environmental controls on phytoplankton community composition in the Thames plume, U.K.

The aim of this study was to investigate controls on the phytoplankton community composition and biogeochemistry of the estuarine plume zone of the River Thames, U.K. using an instrumented moored buoy for in situ measurements and preserved sample collection, and laboratory-based measurements from samples collected at the same site. Instrumentation on the moored buoy enabled high frequency measurements of a suite of environmental variables including in situ chlorophyll, water-column integrated irradiance, macronutrients throughout an annual cycle for 2001 e.g. nitrate and silicate, and phytoplankton biomass and species composition. The Thames plume region acts as a conduit for fluvial nutrients into the wider southern North Sea with typical winter concentrations of 45 μM nitrate, 17 μM silicate and 2 μM phosphate measured. The spring bloom resulted from water-column integrated irradiance increasing above 60 W h m^− 2 d^− 1 and was initially dominated by a diatom bloom mainly composed of Nitzschia sp. and Odontella sinesis. The spring bloom then switched after  30 days to become dominated by the flagellate Phaeocystis reaching a maximum chlorophyll concentration of 37.8 μg L^− 1. During the spring bloom there were high numbers of the heterotrophic dinoflagellates Gyrodinium spirale and Katodinium glaucum that potentially grazed the phytoplankton bloom. This diatom–flagellate switch was predicted to be due to a combination of further increasing water-column integrated irradiance > 100 W h m^− 2 d^− 1 and/or silicate reaching potentially limiting concentrations (< 1 μM). Post spring bloom, diatom dominance of the lower continuous summer phytoplankton biomass occurred despite the low silicate concentrations (Av. 0.7 μM from June–August). Summer diatom dominance, generally due to Guinardia delicatula, was expected to be as a result of microzooplankton grazing, dominated by the heterotrophic dinoflagellate Noctiluca scintillans, controlling 0.7–5.0 μm ‘flagellate’ fraction of the phytoplankton community with grazing rates up to 178% of ‘flagellate’ growth rate. The Thames plume region was therefore shown to be an active region of nutrient and phytoplankton processing and transport to the southern North Sea. The use of a combination of moorings and ship-based sampling was essential in understanding the factors influencing nutrient transport, phytoplankton biomass and species composition in this shelf sea plume region.     

Microzooplankton composition,biomass and grazing rates along the WOCE SR3 line between Tasmania and Antarctica

Microzooplankton species composition and grazing rates on phytoplankton were investigated along a transect between ∼46 and 67°S, and between 140 and 145°E. Experiments were conducted in summer between November 2nd and December 14th in 2001. The structure of the microbial food web changed considerably along the transect and was associated with marked differences in the physical and chemical environment encountered in the different water masses and frontal regions. On average microzooplankton grazing experiments indicated that 91%, 102%, and 157%, (see results) of the phytoplankton production would be grazed in the <200, <20 and <2 μm size fractions, respectively, indicating microzooplankton grazing was potentially constraining phytoplankton populations (<200 μm) along most of the transect. Small ciliates in general and especially oligotrich species declined in importance from the relatively warm, Southern Subtropical Front waters (6.8 μg C/L) to the colder waters of the southern branch of the Polar Front (S-PF), (∼0.5 μg C/L) before increasing again near the Antarctic landmass. Large changes in microzooplankton dominance were observed, with heterotrophic nanoflagellates (HNF), ciliates and larger dinoflagellates having significant biomass in different water masses. HNF were the dominant grazers when chlorophyll a was low in areas such as the Inter-Polar Frontal Zone (IPFZ), while in areas of elevated biomass such as the S-PF and Southern Antarctic Circumpolar Current (SACC), a mix of copepod nauplii and large heterotrophic and mixotrophic dinoflagellates tended to dominate the grazing community. In the S-PF and SACC water masses the tight coupling observed between the microzooplankton grazers and phytoplankton populations over most of the rest of the transect was relaxed. In these regions grazing was low on the >20 μm size fraction of chlorophyll a, which dominated the biomass, while smaller diatoms and nanoplankton in the <20 μm size fraction were still heavily grazed. The lack of grazing pressure on large phytoplankton contributes to this region's potential to export carbon with larger cells known to have higher sinking rates.     

Vertical flux of biogenic matter during a Lagrangian study off the NW Spanish continental margin

Lagrangian experiments with short-term, drifting sediment traps were conducted during a cruise on RRS Charles Darwin to the NW coast of Spain to study the vertical flux and composition of settling biogenic matter. The cruise was split into two legs corresponding to (i) a period of increased production following an upwelling event on the continental shelf (3–10 August 1998) and (ii) an evolution of a cold water filament originating from the upwelled water off the shelf (14–19 August). The export of particulate organic carbon (POC) from the upper layer (0–60m) on the shelf was 90–240mgC.m⁻².d⁻¹ and off the shelf was 60–180mgC.m⁻².d⁻¹. Off shelf the POC flux at 200m was 50–60mg.m⁻².d⁻¹. A modest sedimentation of diatoms (15–30mgC.m⁻².d⁻¹) after the upwelling was associated with increased vertical flux of chlorophyll a (1.8–2.1mg.m⁻².d⁻¹) and a decrease of the POC:PON molar ratio of the settled material from 9 to 6.4. Most of the pico-, nano-, and microplankton in the settled material were flagellates; diatoms were significant during the on shelf and dinoflagellates during the off shelf leg. Off shelf, the exponential attenuation of POC flux indicated a strong retention capacity of the plankton community between 40 and 75m. POC:PON ratio of the settled particulate matter decreased with depth and the relative portion of flagellates increased, suggesting a novel, flagellate and aggregate mediated particulate flux in these waters. Export of POC from the euphotic layer comprised 14–26% of the integrated primary production per day during the on shelf leg and 25–42% during the off shelf leg, which characterises the importance of sedimentation in the organic carbon budget of these waters.     

台湾海峡小型浮游动物的摄食对夏季藻华演替的影响

于2004年8月1~6日对台湾海峡南部近岸的藻华过程进行了定点连续跟踪观测,用稀释法研究了浮游植物的生长率和小型浮游动物对浮游植物的摄食死亡率,同时运用高效液相色谱(HPLC)技术,分析了浮游植物不同光合色素类群的生长率和摄食死亡率.结果表明,观测期间处于藻华的消退期.8月1日时,浮游植物生物量(叶绿素a)和丰度分别为2.04μg/dm3和2.99×105个/dm3,主要优势种为尖刺伪菱形藻(Pseudo-nitzschia pungens)、冰河拟星杆藻(Asterionellopsis glacialis)和中肋骨条藻(Skeletonema costatum),8月6日时,浮游植物生物量和丰度分别减为0.37μg/dm3和1.54×104个/dm3;而蓝藻和甲藻的丰度和比例则呈现出逐渐增加的趋势,所占的比重分别从1日的0.04%和0.85%增加到6日的9.59%和41.97%.小型浮游动物主要由无壳纤毛虫、砂壳纤毛虫、红色中缢虫(Mesodinium rubrum)和异养甲藻等类群组成,总丰度于8月2日达到最大值,为3640个/dm3,之后逐渐减少,6日时,仅为436个/dm3.观测期间,小型浮游动物在群落组成上虽一直以无壳纤毛虫和异养甲藻为主,但在具体的类群结构上却表现出了一定的差异,30μm以下的无壳纤毛虫和异养甲藻总体呈下降的趋势,而红色中缢虫、砂壳纤毛虫和大于50μm的无壳纤毛虫总体呈增加的趋势.观测期间,浮游植物的生长率为0.40~0.91d-1,小型浮游动物的摄食率为0.26~1.34d-1,摄食率和生长率总体呈逐渐下降的趋势.结果还表明,小型浮游动物的摄食率与叶绿素a具有很好的相关性(R2=0.89),对各光合色素类群的现存量和初级生产力均具有较高的摄食压力(分别为37.97%~82.24%和70.71%~281.33%),是藻华消亡的重要原因之一;此外,小型浮游动物对甲藻和蓝藻的避食行为,可能是观测期间由“硅藻”水华向“硅藻-甲藻”水华转变的重要原因之一.     

Plankton community diversity from bacteria to copepods in bloom and non-bloom conditions in the Celtic Sea in spring

The plankton community composition comprising heterotrophic bacteria, pro-/eukaryotes, heterotrophic nanoflagellates, microzooplankton and mesozooplankton was assessed during the spring bloom and at non-bloom stations in the English Channel and Celtic Sea between 6 and 12 April 2002. Non-bloom sites were characterised by a dominance of pro-/eukaryotic phytoplankton <20 μm, higher abundance of heterotrophic nanoflagellates, microzooplankton standing stocks ranging between 60 and 380 mg C m⁻², lower mesozooplankton diversity and copepod abundance of between 760 and 2600 ind m⁻³. Within the bloom, the phytoplankton community was typically dominated by larger cells with low abundance of pro-/eukaryotes. Heterotrophic nanoflagellate cell bio-volume decreased leading to a reduction in biomass whereas microzooplankton biomass increased (360–1500 mg C m⁻²) due to an increase in cell bio-volume and copepod abundance ranged between 1400 and 3800 ind m⁻³. Mesozooplankton diversity increased with an increase in productivity. Relationships between the plankton community and environmental data were examined using multivariate statistics and these highlighted significant differences in the abiotic variables, the pro-/eukaryotic phytoplankton communities, heterotrophic nanoflagellate, microzooplankton and total zooplankton communities between the bloom and non-bloom sites. The variables which best described variation in the microzooplankton community were temperature and silicate. The spatial variation in zooplankton diversity was best explained by temperature. This study provides an insight into the changes that occur between trophic levels within the plankton in response to the spring bloom in this area.     

Seasonality of microzooplankton grazing in the northern Wadden Sea

A sequence of nine dilution experiments was conducted according to Landry and Hassett [Landry, M.R., Hassett, R.P., 1982. Estimating the grazing impact of marine microzooplankton. Mar. Biol. 67, 283–288] in the northern Wadden Sea from March until October 2004 to investigate the seasonality of microzooplankton grazing. From March until April, no grazing was observed. Microzooplankton grazing started in May (0.66 d^− 1) and increased until August (1.22 d^− 1). In October microzooplankton grazing was low again (0.17 d^− 1). Phytoplankton growth rates varied between 0 and 1.1 d^− 1. Since the reliability of dilution experiments is still frequently discussed in literature, we tested if our data obtained by dilution experiments reflected short-term in situ phytoplankton dynamics of the study site. We scaled experimental growth rates to water column irradiance, calculated short-term chlorophyll-a dynamics and compared the results to in situ measured chlorophyll-a concentrations. Calculated chlorophyll-a concentrations correlated significantly with in situ measured chlorophyll-a concentrations but slightly overestimated the in situ measured chlorophyll-a. This overestimation was in the range of phytoplankton assimilation reported for the Wadden Sea benthos. We will show that microzooplankton grazing had a large impact during the Phaeocystis bloom and during summer suggesting that a large proportion of phytoplankton biomass remained the pelagic food web. Microzooplankton grazing did not impact the diatom spring bloom and its demise.     

Contrasts in sedimentation flux below the southern California Current in late 1996 and during the El Niño event of 1997–1998

The vertical flux of particulate matter at 330 m depth in San Lázaro Basin off southern Baja California ranged from 63 to 587 mg m⁻² d⁻¹ between August and November 1996. Organic carbon contents were between 5.6 and 14.8%, yielding flux rates of 9–40 mgC m⁻² d⁻¹. In December 1997 and January 1998, at the height of the strong El Niño event, the respective fluxes (47–202 mg m⁻² d⁻¹ and 3–8 mgC m⁻² d⁻¹) were comparable. The February–June 1998 records, however, revealed sharply reduced mass (1–6 mg m⁻² d⁻¹) and organic carbon (0.2–0.8 mgC m⁻² d⁻¹) fluxes. The organics collected in 1996 were predominantly autochthonous (δ¹³C=−22‰; C/N=8). The variations in δ¹⁵N (8.3–11.0‰) suggest an alternation of new and regenerated production, possibly associated with fluctuations in the intensity of deep mixing during that autumn. The relatively high organic matter fluxes in December 1997 appear to be associated with regenerated production. The average composition from February to June 1998 (δ¹³C=−23.6‰; ¹⁵N=11.7‰; C/N=10.5) indicates degraded material of marine origin. The maximum δ¹⁵N value found (14‰) suggests that deeper, denitrified waters were brought to the surface and possibly advected laterally. Regime changes in the waters of the basin occur at 6–10 week intervals, evidenced by concurrent shifts in most of the measured parameters, including fecal pellet types and metal chemistry. The marine snow-dominated detritus collected showed a shift from a mixed diatom-rich-radiolarian-coccolith assemblage in late 1996 to a coccolith-dominated assemblage, including the contents of fecal pellets, during the 1997–1998 El-Niño period. T–S profiles, plankton analysis and chlorophyll contents of the upper water column indicated that the strong phytoplankton bloom, normally associated with seasonal upwelling along the Pacific coast of Baja, did not occur during the spring of 1998. The persistence of oligotrophic conditions during the 1997–1998 El Niño event favored the dominance of nanoplankton and reduced the vertical flux of particles.     

Microzooplankton grazing impact in the Western Arctic Ocean

Microzooplankton grazing impact on phytoplankton was assessed using the Landry–Hassett dilution technique in the Western Arctic Ocean during spring and summer 2002 and 2004. Forty experiments were completed in a region encompassing productive shelf regions of the Chukchi Sea, mesotrophic slope regions of the Beaufort Sea off the North Slope of Alaska, and oligotrophic deep-water sites in the Canada Basin. A variety of conditions were encountered, from heavy sea-ice cover during both spring cruises, moderate sea-ice cover during summer of 2002, and light to no sea ice during summer of 2004, with a concomitant range of trophic conditions, from low chlorophyll-a (Chl-a; <0.5 μg L⁻¹) during heavy ice cover in spring and in the open basin, to late spring and summer shelf and slope open-water diatom blooms with Chl-a >5 μg L⁻¹. The microzooplankton community was dominated by large naked ciliates and heterotrophic gymnodinoid dinoflagellates. Significant, but low, rates of microzooplankton herbivory were found in half of the experiments. The maximum grazing rate was 0.16 d⁻¹ and average grazing rate, including experiments with no significant grazing, was 0.04±0.06 d⁻¹. Phytoplankton intrinsic growth rates varied from the highest values of about 0.4 d⁻¹ to the lowest values of zero to slightly negative growth, on average 0.16±0.15 d⁻¹. Light limitation in spring and post-bloom senescence during summer were likely explanations of observed low phytoplankton growth rates. Microzooplankton grazing consumed 0–120% (average 22±26%) of phytoplankton daily growth. Grazing and growth rates found in this study were low compared to rates reported in another Arctic system, the Barents Sea, and in major geographic regions of the world ocean.     

Microzooplankton grazing and selective feeding during bloom periods in the Tolo Harbour area as revealed by HPLC pigment analysis

Dilution experiments were conducted to investigate microzooplankton grazing impact on phytoplankton of different taxonomic groups and size fractions (< 5, 5–20, 20–200 μm) during spring and summer bloom periods at two different sites (inner Tolo Harbour and Tolo Channel) in the Tolo Harbour area, the northeastern coastal area of Hong Kong. Experiments combined with HPLC pigment analysis in three phytoplankton size fractions measured pigment and size specific phytoplankton growth rates and microzooplankton grazing rates. Pigment-specific phytoplankton growth rates ranged between 0.08 and 3.53 d^− 1, while specific grazing rates of microzooplankton ranged between 0.07 and 2.82 d^− 1. Highest specific rates of phytoplankton growth and microzooplankton grazing were both measured in fucoxanthin in 5–20 μm size fraction in inner Tolo Harbour in summer, which coincided with the occurrence of diatom bloom. Results showed significant correlations between phytoplankton growth and microzooplankton grazing rates. Microzooplankton placed high grazing pressure on phytoplankton community. High microzooplankton grazing impact on alloxanthin (2.63–5.13) suggested strong selection toward cryptophytes. Our results provided no evidence for size selective grazing on phytoplankton by microzooplankton.     

Community structure and grazing of the nano-microzooplankton on the continental shelf of the Bay of Biscay

In order to investigate the parameters controlling the heterotrophic protists (nano-microzooplankton) on the continental shelf of the southern Bay of Biscay, plankton communities and their physico-chemical environment were studied 4 times in February, April, June and September–October 2004 at three stations in the euphotic zone in the Bay of Biscay. The abundance and carbon biomass of heterotrophic protists (ciliates, heterotrophic dinoflagellates and nanoflagellates) as well as all the others groups of plankton (picoplankton, nanophytoplankton, diatoms, autotrophic dinoflagellates, metazoan microzooplankton and mesozooplankton), the environmental parameters and the primary and bacteria production were evaluated at each sampling period. Microzooplankton grazing experiments were undertaken at the same time. Ciliates and heterotrophic dinoflagellates accounted for the main major component of nano- and microzooplankton communities in term of biomass. The total carbon biomass of heterotrophic protists was highest in spring and lowest at the end of summer. The development of heterotrophic protists started after a winter microphytoplankton bloom (principally large diatoms), the biomass was lower in June and was low in September (through inappropriate prey). The carbon requirement of microzooplankton ranged from 50 to more than 100% of daily primary, bacterial and nanoflagellate production. The heterotrophic protist community was predominantly constrained by bottom-up control in spring and at the end of summer via food availability and quality.     

Distribution and sinking rates of phytoplankton, detritus, and particulate biogenic silica in the Laptev Sea and Lena River (Arctic Siberia)

Concentrations and sinking rates of particulate biogenic silica (BSi), chlorophyll a (chl a) and phaeopigments (phae) (< 3 μm, 3–10 μm, > 10 μm and total), as well as the abundances of the major phytoplankton species, were studied during September 1991 in the Eastern Laptev Sea and the lower Lena River (Siberian Arctic). The highest chl a concentrations were found in two major “new” production regimes of the study area: (1) a deep chl a maximum (5.8 mg chl a m⁻³) (formed by the diatom Chaetoceros socialis) at 30 m depth on the outer shelf of the northern Laptev Sea, and (2) in the Lena River, where the phytoplankton community was dominated by fresh water diatoms (1.5 to 4.5 mg chl a m⁻³). Elevated chl a concentrations were also found in the river plume phytoplankton community (dominated by brackish water diatoms), NE of the Lena delta. In the Laptev Sea, the low chl a (0.1 to 3 mg chl a m⁻³) and high phae concentrations (0.5 to 14 mg phae m⁻³) indicated that the phytoplankton community (dominated by picoplanktic algae and nanoflagellates) was already senescent and affected by grazing losses. Biogenic silica values were highest in the Lena River (4 to 17 μM) as compared to the low values found in the Laptev Sea (0.3 to 4 μM). The large chl a size fraction, phae and BSi in the Lena River samples revealed the highest measured sinking rates (1.4, 2.3, and 1.5 m d⁻¹, respectively). The formation of a strong halocline, decreasing turbulence, and possible nutrient deficiency resulted in death, disintegration and rapid sedimentation of fresh water diatoms. This was accompanied by a decrease in the BSi concentration and growth of the picoplanktic size fraction (< 3 μm) in the estuarine mixing zone (Gulf of Buorkhaya). Only a minor part of BSi was bound to intact diatom cells (< 3%) in the surface layer, most of which being apparently associated with detrital particles. In the Lena River, approximately 12% of the total silica was bound to BSi fraction, yet elsewhere in the Laptev Sea and in the estuarine mixing zone the BSi:total silica ratio was ≤ 5%. Thus, the results reflected the successional stage of a late summer phytoplankton community, characterized by dominance of small autotrophs and patchy distribution of senescent diatoms no longer able to affect the relative high levels of dissolved silica supplied by the Lena River.     

Photosynthate allocation in a temperate sea over an annual cycle: the relationship between protein synthesis and phytoplankton physiological state

The seasonal and vertical variations in the patterns of photosynthate allocation into biomolecules by natural phytoplankton assemblages were determined, together with their species composition, in a coastal station of the central Cantabrian Sea (southern Bay of Biscay). Chlorophyll-a concentration ranged from values below 20 mg m⁻² in winter to values above 80 mg m⁻² during spring and during an upwelling event in summer. Low primary production rates (<300 mgC m⁻² d^-1) were measured during winter and during summer stratification periods. The rate of C fixation during summer upwelling conditions exceeded 3500 mgC m⁻² d⁻¹. In terms of photosynthate partitioning, proteins were the dominant fraction, as they typically accounted for >30% of total photo-assimilated C, with polysaccharides and low molecular weight metabolites showing incorporation percentages around 10–30%. Relative C incorporation into lipids was generally <15%. Recurrent patterns of vertical variability in photosynthate partitioning were observed: the relative synthesis of proteins increased toward the bottom of the euphotic zone, whereas the relative C incorporation into polysaccharides and lipids tended to be higher near the surface. When primary production decreased, the synthesis of proteins was maintained more than that of other molecules. Throughout the year, the relative synthesis of proteins was inversely correlated with phytoplankton biomass, production and growth rate. The conservation of protein synthesis under growth-limiting conditions and the enhancement of lipid and polysaccharide synthesis when irradiance is high seem to constitute general patterns of photosynthate partitioning in marine phytoplankton. In our study, these patterns represented metabolic strategies of phytoplankton in response to changing environmental factors, rather than the effect of variations in the species composition of the community.     

Microzooplankton herbivory in the Ross Sea, Antarctica

Microplankton abundances and phytoplankton mortality rates were determined at six stations during four cruises spanning three seasons in the Ross Sea polynya, Antarctica (early spring, Oct.–Nov. 1996; mid-late summer, Jan.–Feb. 1997; fall, Apr. 1997; mid-late spring, Nov.–Dec. 1997). Rates of microzooplankton herbivory were measured using a modified dilution technique, as well as by examining the rate of disappearance of phytoplankton (chlorophyll) in samples incubated in the dark (i.e. grazing in the absence of phytoplankton growth). Strong seasonal cycles of phytoplankton and microzooplankton abundance were observed during the study. Microzooplankton abundance varied by more than three orders of magnitude during the four cruises, and was positively correlated with phytoplankton biomass over the entire data set. Nevertheless, microzooplankton grazing was insufficient to impact significantly phytoplankton standing stocks during most of the experiments performed in this perenially cold environment. Only thirteen out of a total of 51 experiments yielded phytoplankton mortality rates that were significantly different from zero. The highest mortality rate observed in this study (0.26 d⁻¹) was modest compared with maximal rates that have been observed in temperate and tropical ecosystems. Results from twenty experiments examining the rate of decrease of phytoplankton biomass during incubations in the dark agreed quite well with the results of the dilution experiments performed at the same time. The range of mortality rates for the dark incubations was −0.09–0.06 d⁻¹, and the average was essentially zero (−0.01 d⁻¹). That is, chlorophyll concentration was virtually unchanged in samples incubated in the dark for 3 d. A number of factors appeared to contribute to the very low rates of microbial herbivory observed, including low water temperature, and the size and taxonomic composition of the phytoplankton assemblage. Based on our results we conclude that the seasonal, massive phytoplankton blooms observed in the Ross Sea are due, in part, to low rates of removal by microbial herbivores.