Trophic coupling between <Emphasis Type="Italic">Synechococcus</Emphasis> and pigmented nanoflagellates in the coastal waters of Taiwan,western subtropical Pacific

In our attempt to characterize the interaction of trophic coupling between Synechococcus and pigmented nanoflagellates (PNFs), successive size-fraction experiments were performed at a coastal station on the northeast coast of Taiwan from June, 2005 to January, 2006. By estimating the growth rate and grazing rate of Synechococcus in the presence of nanoflagellates of different sizes, we truncated the food web by removing organisms with different body sizes (<2 μm, <5 μm, <10 μm, and <20 μm). The growth rates of Synechococcus ranged from −0.016 to 0.051 h⁻¹ during the experimental period, suggesting that temperature was a primary mechanism controlling Synechococcus growth. In addition to size and relative biomass of pigmented nanoflagellates and Synechococcus, it is suggested that community structures played an important role in trophic link. Furthermore, we conclude that the trophic cascading effect in the northeast coast of Taiwan includes: 1) high grazing rates at night in the warm season; 2) the Synechococcus biomass generally exceeds the grazing threshold (6 × 10⁴ cells mL⁻¹); and 3) the biomass ratio of <5 μm PNFs to >5 μm PNFs should be 1:1 to 2:1.     

Heterotrophic bacterial and <Emphasis Type="Italic">Synechococcus</Emphasis> spp. Growth and mortality along the inshore-offshore in the East China Sea in summer

This study used the dilution method to examine growth and grazing rates of heterotrophic bacteria and an autotrophic picoplankton, Synechococcus spp., from 1 to 11 July 2007 in the East China Sea. The main influence of oceanographic conditions in this aquatic system was the introduction of fresh, high-nutrient water from Changjiang River and the extremely nutrient-poor, high-salinity waters of Kuroshio Water. In these experiments, deviation from linearity in the relationship between dilution factor and net growth rate was significant in a large number of cases. Growth rates for heterotrophic bacteria ranged from 0.024 to 0.24, and for Synechococcus spp. from 0.03 to 0.21 h⁻¹. Grazing rates ranged from 0.02 to 0.19 and 0.01 to 0.13 h⁻¹, respectively. The spatial variations of Synechococcus spp. production to the primary production ratio (SP/PP) were low (<5%) in high Chl a environments and increased exponentially in low Chl a environments, indicating that Synechococcus spp. contributes to a large extent to the photosynthetic biomass in the open sea, especially in the more oligotrophic Kuroshio Water. Furthermore, the results of our dilution experiments suggest that nanoflagellates largely depend on heterotrophic bacteria as an important energy source. On average, heterotrophic bacteria contributes to 76 and 59% of carbon consumed by nanoflagellates within the plume (salinity <31) and outside of it (salinity >31).     

Phytoplankton growth and microzooplankton grazing in the Sea of Okhotsk during late summer of 2006

The Sea of Okhotsk is one of the most productive marine basins in the world ocean and plays an important role in transport of organic carbon and iron to the western subarctic Pacific. We report the first measurements of phytoplankton growth and microzooplankton grazing rates in the Sea of Okhotsk, in late summer of 2006. The study area can be divided into two areas: nutrient-sufficient waters on the continental shelf along the east coast of Sakhalin Island and in the vicinity of Bussol Strait, and surface nutrient-depleted waters beyond the shelf break and in the vicinity of Sakhalin Bay. Phytoplankton growth rate in the studied area was strongly affected by nutrient availability, with high phytoplankton growth rate (0.55±0.14 d^?1) in the nutrient-replete region and severely depressed growth (0.03±0.05 d^?1) in the nutrient-depleted region. On the other hand, microzooplankton grazing rates in both the nutrient-replete and nutrient-depleted regions were approximately the same (0.26±0.20 d^?1 vs. 0.27±0.24 d^?1). Consequently, microzooplankton grazing consumed <50% of the phytoplankton growth in nutrient-rich waters but >3 times the phytoplankton growth in nutrient-depleted waters. Phytoplankton physiological condition as measured by the maximum photochemical quantum efficiency (F_v/F_m) of algal photosystem II (PS II) showed a general trend in agreement with the in situ growth rate of phytoplankton. In contrast to the phytoplankton community, picophytoplankton, especially the cyanobacteria Synechococcus, showed no nutrient effect on their growth, and the growth and mortality rates were well balanced, suggesting that they have a low nutrient requirement and their biomass was controlled principally by microzooplankton grazing.     

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.     

Carbon fluxes through major phytoplankton groups during the spring bloom and post-bloom in the Northwestern Mediterranean Sea

The carbon flux through major phytoplankton groups, defined by their pigment markers, was estimated in two contrasting conditions of the Northwestern Mediterranean open ocean ecosystem: the spring bloom and post-bloom situations (hereafter Bloom and Post-bloom, respectively). During Bloom, surface chlorophyll a (Chl a) concentration was higher and dominated by diatoms (53% of Chl a), while during Post-bloom Synechococcus (42%) and Prymnesiophyceae (29%) became dominant. The seawater dilution technique, coupled to high pressure liquid chromatography (HPLC) analysis of pigments and flow cytometry (FCM), was used to estimate growth and grazing rates of major phytoplankton groups in surface waters. Estimated growth rates were corrected for photoacclimation based on FCM-detected changes in red fluorescence per cell. Given the 30% average decrease in the pigment content per cell between the beginning and the end of the incubations, overlooking photoacclimation would have resulted in a 0.40 d^?1 underestimation of phytoplankton growth rates. Corrected average growth rates (μ_o) were 0.90±0.20 (SD) and 0.40±0.14 d^?1 for Bloom and Post-bloom phytoplankton, respectively. Diatoms, Cryptophyceae and Synechococcus were identified as fast-growing groups and Prymnesiophyceae and Prasinophyceae as slow-growing groups across Bloom and Post-bloom conditions. The higher growth rate during Bloom was due to dominance of phytoplankton groups with higher growth rates than those dominating in Post-bloom. Average grazing rates (m) were 0.58±0.20 d^?1 (SD) and 0.31±0.07 d^?1. The proportion of phytoplankton growth consumed by microzooplankton grazing (m/μ_o) tended to be lower in Bloom (0.69±0.34) than in Post-bloom (0.80±0.08). The intensity of nutrient limitation experienced by phytoplankton indicated by μ_o/μ_n (where μ_n is the nutrient-amended growth rate), was similar during Bloom (0.78) and Post-bloom (0.73). Primary production from surface water (PP) was estimated with ¹⁴C incubations. A combination of PP and Chl a synthesis rate yielded C/Chl a ratios of 34±21 and 168±75 (g:g) for Bloom and Post-bloom, respectively. Transformation of group-specific Chl a fluxes into carbon equivalents confirmed the dominant role of diatoms during Bloom and Synechococcus and Prymnesiophyceae during Post-bloom.     

Picophytoplankton dynamics and production in the Arabian Sea during the 1995 Southwest Monsoon

Phytoplankton community structure is expected to shift to larger cells (e.g., diatoms) with monsoonal forcing in the Arabian Sea, but recent studies suggest that small primary producers remain active and important, even in areas strongly influenced by coastal upwelling. To better understand the role of smaller phytoplankton in such systems, we investigated growth and grazing rates of picophytoplankton populations and their contributions to phytoplankton community biomass and primary productivity during the 1995 Southwest Monsoon (August–September). Environmental conditions at six study stations varied broadly from open-ocean oligotrophic to coastal eutrophic, with mixed-layer nitrate and chlorophyll concentrations ranging from 0.01 to 11.5 μM NO₃ and 0.16 to 1.5 μg Chl a. Picophytoplankton comprised up to 92% of phytoplankton carbon at the oceanic stations, 35% in the diatom-dominated coastal zone, and 26% in a declining Phaeocystis bloom. Concurrent in situ dilution and ¹⁴C-uptake experiments gave comparable ranges of community growth rates (0.53–1.05 d⁻¹ and 0.44–1.17 d⁻¹, to the 1% light level), but uncertainties in C:Chl a confounded agreement at individual stations. Microzooplankton grazing utilized 81% of community phytoplankton growth at the oligotrophic stations and 54% at high-nutrient coastal stations. Prochlorococcus (PRO) was present at two oligotrophic stations, where its maximum growth approached 1.4 d⁻¹ (two doublings per day) and depth-integrated growth varied from 0.2 to 0.8 d⁻¹. Synechococcus (SYN) growth ranged from 0.5 to 1.1 d⁻¹ at offshore stations and 0.6 to 0.7 d⁻¹ at coastal sites. Except for the most oligotrophic stations, growth rates of picoeukaryotic algae (PEUK) exceeded PRO and SYN, reaching 1.3 d⁻¹ offshore and decreasing to 0.8 d⁻¹ at the most coastal station. Microzooplankton grazing impact averaged 90, 70, and 86% of growth for PRO, SYN, and PEUK, respectively. Picoplankton as a group accounted for 64% of estimated gross carbon production for all stations, and 50% at high-nutrient, upwelling stations. Prokaryotes (PRO and SYN) contributed disproportionately to production relative to biomass at the most oligotrophic station, while PEUK were more important at the coastal stations. Even during intense monsoonal forcing in the Arabian Sea, picoeukaryotic algae appear to account for a large portion of primary production in the coastal upwelling regions, supporting an active community of protistan grazers and a high rate of carbon cycling in these areas.     

Abundance, Biomass, Production and Trophic Roles of Micro- and Net-Zooplankton in Ise Bay, Central Japan, in Winter

We investigated the geographical variations in abundance and biomass of the major taxonomic groups of micro- and net-zooplankton along a transect through Ise Bay, central Japan, and neighboring Pacific Ocean in February 1995. The results were used to estimate their secondary and tertiary production rates and assess their trophic roles in this eutrophic embayment in winter. Ise Bay nourished a much higher biomass of both micro- and net-zooplankton (mean: 3.79 and 13.9 mg C m^–3, respectively) than the offshore area (mean: 0.76 and 4.47 mg C m^–3, respectively). In the bay, tintinnid ciliates, naked ciliates and copepod nauplii accounted for, on average, 69, 18 and 13% of the microzooplankton biomass, respectively. Of net-zooplankton biomass, copepods (i.e. Acartia, Calanus, Centropages, Microsetella and Paracalanus) formed the majority (mean: 63%). Average secondary production rates of micro- and net-zooplankton in the bay were 1.19 and 1.87 mg C m^–3d^–1 (or 23.1 and 36.4 mg C m^–2d^–1), respectively, and average tertiary production rate of net-zooplankton was 0.75 mg C m^–3d^–1 (or 14.6 mg C m^–2d^–1). Available data approximated average phytoplankton primary production rate as 1000 mg C m^–2d^–1 during our study period. The transfer efficiency from primary production to zooplankton secondary production was 6.0%, and the efficiency from secondary production to tertiary production was 25%. The amount of food required to support the zooplankton secondary production corresponded to 18% of the phytoplankton primary production or only 1.7% of the phytoplankton biomass, demonstrating that the grazing impact of herbivorous zooplankton was minor in Ise Bay in winter.     

Top-down control of spring surface phytoplankton blooms by microzooplankton in the Central Yellow Sea,China

Phytoplankton growth and microzooplankton grazing were studied during the 2007 spring bloom in Central Yellow Sea. The surveyed stations were divided to pre-bloom phase (Chl a concentration less than 2 μg L⁻¹), and bloom phase (Chl a concentration greater than 2 μg L⁻¹). Shipboard dilution incubation experiments were carried out at 19 stations to determine the phytoplankton specific growth rates and the specific grazing rates of microzooplankton on phytoplankton. Diatoms dominated in the phytoplankton community in surface waters at most stations. For microzooplankton, Myrionecta rubra and tintinnids were dominant, and heterotrophic dinoflagellate was also important in the community. Phytoplankton-specific growth rates, with an average of 0.60±0.19 d⁻¹, were higher at pre-bloom stations (average 0.62±0.17 d⁻¹), and lower at the bloom stations (average 0.59±0.21 d⁻¹), but the difference of growth rates between bloom and pre-bloom stations was not statistically significant (t test, p=0.77). The phytoplankton mortality rate by microzooplankton grazing averaged 0.41±0.23 d⁻¹ at pre-bloom stations, and 0.58±0.31 d⁻¹ during the blooms. In contrast to the growth rates, the statistic difference of grazing rates between bloom and pre-bloom stations was significant (after removal of outliers, t test, p=0.04), indicating the importance of the top-down control in the phytoplankton bloom processes. Average potential grazing efficiency on primary productivity was 66% at pre-bloom stations and 98% at bloom stations, respectively. Based on our results, the biomass maximum phase (bloom phase) was not the maximum growth rate phase. Both phytoplankton specific growth rate and net growth rate were higher in the pre-bloom phase than during the bloom phase. Microzooplankton grazing mortality rate was positively correlated with phytoplankton growth rate during both phases, but growth and grazing were highly coupled during the booming phase. There was no correlation between phytoplankton growth rate and cell size during the blooms, but they were positive correlated during the pre-bloom phase. Our results indicate that microzooplankton grazing is an important process controlling the growth of phytoplankton in spring bloom period in the Central Yellow Sea, particularly in the “blooming” phase.     

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.     

Ingestion rates and grazing impact of the brackwater mussel Brachidontes virgiliae in Lake St Lucia,iSimangaliso Wetland Park,South Africa

Bivalves feed on a combination of phytoplankton and zooplankton and have the potential to impact considerably the planktonic biomass, especially when they occur in high densities, such as in oyster and mussel beds. The brackwater mussel Brachidontes virgiliae is numerically dominant during wet phases within Africa’s largest estuarine lake, St Lucia, in the iSimangaliso Wetland Park on the east coast of South Africa. The ingestion rates and potential grazing impact of this small mussel (maximum shell length = 2.5 cm) were estimated for both the wet and dry seasons using an in situ gut fluorescence technique. Ingestion rates were higher during the wet season (5.78 µg pigment ind.^?1 d^?1) than during the dry season (4.44 µg pigment ind.^?1 d^?1). This might be explained by the increased water temperature and food availability during the wet season. Because of the patchy distribution of mussel populations, there could be higher localised grazing impact near mussel aggregations. Results showed a potential grazing impact of up to 20 times the available phytoplankton biomass at specific sites. These high grazing impacts have the potential to deplete phytoplankton stocks in the lake, especially during wet phases in the northern reaches, where mussel densities are highest. This needs to be factored into ecological models of Lake St Lucia, because the system might function differently during increased flood events.     

Distribution and contribution of major phytoplankton groups to carbon cycling across contrasting conditions of the subtropical northeast Atlantic Ocean

The relation between trophic regime and phytoplankton composition and function in oceanic systems is well accepted in oceanography. However, the relative dynamics and carbon cycling contributions of different phytoplankton groups across gradients of ocean richness are not fully understood. In this work we investigated phytoplankton dynamics along two transects from the NW African coastal upwelling to open-ocean waters of the north Atlantic subtropical gyre. We adopted a pigment-based approach to characterize community structure and to quantify group-specific growth and grazing rates and associated carbon fluxes. Changes in pigment cell concentration during the incubation experiments due to photoadaptation were corrected to obtain reliable rates. The oceanic region was dominated by Prochlorococcus (PRO) (45±7% of total chlorophyll a) while diatoms dominated in upwelling waters (40±37%). Phytoplankton grew faster (μ=0.78±0.26 d⁻¹) and free of nutrient limitation (μ/μ_n=0.98±0.42) in the coastal upwelling region, with all groups growing at similar rates. In oceanic waters, the growth rate of bulk phytoplankton was lower (μ=0.52±0.16 d⁻¹) and nutrient limited (μ/μ_n=0.68±0.19 d⁻¹). Diatoms (0.80±0.39 d⁻¹) and Synechococcus (SYN) (0.72±0.25 d⁻¹) grew faster than Prymnesiophyceae (PRYMN) (0.62±0.26 d⁻¹) and PRO (0.46±0.18 d⁻¹). The growth rates of PRO and SYN were moderately nutrient limited (μ/μ_n=0.81 and 0.91, respectively), while the limitation for diatoms (μ/μ_n=0.71) and PRYMN (μ/μ_n=0.37) was more severe. Microzooplankton grazing rate was higher in upwelling (0.68±0.32 d⁻¹) than in oceanic waters (0.37±0.19 d⁻¹), but represented the main loss pathway for phytoplankton in both systems (m/μ=0.90±0.32 and 0.69±0.24, respectively). Carbon flux through phytoplankton, produced and grazed, increased from offshore to coastal (∼2 to ∼200 μg C L⁻¹ d⁻¹), with diatoms dominating the flux in the upwelling region (52%) while PRYMN (40%) and PRO (30%) dominated in the open ocean.     

Phytoplankton growth and microzooplankton grazing in the subarctic Pacific Ocean and the Bering Sea during summer 1999

Phytoplankton growth and microzooplankton grazing rates were measured by the dilution technique in the subarctic North Pacific Ocean along a west–east transect during summer 1999. Average phytoplankton growth rates without added nutrients (μ₀) were 0.33, 0.41, 0.20 and 0.49 d⁻¹ for the four regions sampled: the Western Gyre, the Bering Sea, the Gulf of Alaska gyre and stations along the Aleutian Trench. Average grazing mortality rates (m) were 0.34, 0.27, 0.20 and 0.49 d⁻¹. Limitation of phytoplankton growth by macronutrients, such as NO₃ and SiO₂, was identified only at a few stations, with overall μ₀/μ_n (μ_n is nutrient-enhanced growth rate) averaging 0.9. Phytoplankton growth and microzooplankton grazing were approximately balanced, as indicated by high m/μ₀ ratio, except in the Bering Sea, where the m/μ₀ ratio was 0.65, indicating the relative importance of the diatom-macrozooplankton grazing food chain and possible higher export flux to the deep layer. Flow cytometric analysis revealed that the growth rates of picoplankton (Synechococcus and picoeukaryotes) were usually much lower than the total phytoplankton community growth rates estimated from chlorophyll a, except for stations in the Gulf of Alaska Gyre, where the growth rates for different populations were about the same. Lower than community-average growth rate for picoplankton indicates larger phytoplankters, presumably diatoms, were growing at a much faster rate. Suppressed phytoplankton growth in the Gulf of Alaska was probably a result of iron limitation.     

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 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.     

Zooplankton herbivory in the Arabian Sea during and after the SW monsoon, 1994

Microzooplankton herbivory in the Arabian Sea was measured using dilution experiments towards the end of the SW monsoon in September and during the intermonsoon to NE monsoon period in November–December 1994. Microzooplankton grazing resulted in a turnover of phytoplankton stocks that ranged from 11 to 49% per day. This was equivalent to grazing fluxes of between 1 and 17 mg C m^-3 d^-1. Depth-integrated microzooplankton herbivory ranged between 161 and 415 mg C m^-2 d^-1 during the SW monsoon cruise, and between 110 and 407 mg C m^-2 d^-1 during the intermonsoon period. Microzooplankton grazed between 4 and 60% of daily primary production, with higher percentages found during the intermonsoon season. Phytoplankton growth rates during the SW monsoon ranged from 0.3 to 1.8 d^-1, with lower values in upwelling waters and higher values in downwelling and oligotrophic areas. During the intermonsoon period, phytoplankton growth was more uniform across the basin and averaged 0.68±0.15 d^-1. Microzooplankton abundance in experimental samples varied between 2800 and 16 162 cells l^-1, equivalent to a biomass of between 1.1 and 7.2 mg C m^-3. The mean cell carbon content of microzooplankton was similar in both periods and ranged from 0.33 to 0.55 ng C cell^-1. Microzooplankton were smallest in downwelling waters and largest in oligotrophic waters. Average clearance rates in those taxa that took up fluorescently-labelled algae ranged from 0.2 to 14 μl ind^-1 hr^-1. Average mesozooplankton grazing rates, derived from biomass data, varied from 19 to 92 mg C m^-2 d^-1; these rates accounted for removal of between 4 and 12% of the daily primary production. Mesozooplankton herbivory was most pronounced in upwelling and downwelling waters and reduced in stratified oligotrophic waters during the SW monsoon period. Microzooplankton herbivory was greater than the average mesozooplankton herbivory at all stations, during both the SW monsoon and intermonsoon periods.     

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.     

Phytoplankton growth and mortality during the 1995 Northeast Monsoon and Spring Intermonsoon in the Arabian Sea

Phytoplankton growth rates and mortality rates were experimentally examined at eight stations in the Arabian Sea along the U.S. JGOFS cruise track during the 1995 Northeast Monsoon (January) and Spring Intermonsoon (March–April). Instantaneous growth rates averaged over an entire cruise were approximately twice as high during the NE Monsoon than during the Spring Intermonsoon period (overall averages of 0.84±0.29 (s.d.) versus 0.44±0.19 d⁻¹). Average herbivore grazing (mortality) rates, however, were quite similar for the two seasons (overall averages of 0.35±0.18 and 0.30±0.17 d⁻¹ for the NE Monsoon and Spring Intermonsoon, respectively). The absolute amounts of phytoplankton biomass consumed during each season also were similar (29 and 25% of standing stock consumed d⁻¹ for the January and March–April cruises, respectively), as were the geographical trends of this removal. These seasonal trends in growth and removal rates resulted in net phytoplankton growth rates that were considerably higher during the January cruise (0.48 d⁻¹) than during the March–April cruise (0.14 d⁻¹). That is, phytoplankton production was more closely balanced during the Spring Intermonsoon season (87% of daily primary production consumed) relative to the NE Monsoon season (49% of daily primary production consumed). Station-to-station variability was high for rate measurements during either cruise. Nevertheless, there was a clear onshore–offshore trend in the absolute rate of removal of phytoplankton biomass (μg chlorophyll consumed l⁻¹ d⁻¹) during both cruises. Coastal stations had removal rates that were typically 2–4 times higher than removal rates at oceanic stations.     

黄海春季水华过程中微微型浮游生物的变化特点

于2007年3—4月在黄海中部海域采用流式细胞术研究了春季水华过程中聚球藻、微微型真核浮游生物和异养细菌的生物量变化。聚球藻和微微真核型浮游生物的生物量与叶绿素a浓度变化基本呈现相反的趋势,在水华前期较高,水华期迅速下降,直至水华后期又有所升高。异养细菌在整个水华过程中变化较小,生物量在水华期最高,与水柱叶绿素a浓度呈极显著正相关(r=0.319,p<0.01)。水华期这三类微微型浮游生物对浮游植物总碳生物量的贡献很低。纤毛虫和鞭毛虫捕食可能是导致聚球藻和微微型真核浮游生物在水华期生物量降低的主要原因。     

河北沿岸微微型浮游植物的分布特征

于2006年7月～ 2007年10月间,分4个季度调查了河北省沿岸微微型浮游植物的丰度和生物量及对浮游植物总生物量的贡献.结果显示:河北近岸海域聚球藻蓝细菌丰度为4.46×103个/mL(0.79×103～ 16.19×103个/mL),生物量(以碳计,下同)为1.31 mg/m3 (0.84～17.47 mg/m3),季节分布特征为秋季＞冬季＞夏季＞春季.微微型光合真核生物丰度为4.43×102个/mL (0.84×102～ 17.47×102个/mL),生物量为1.11mg /m3 (0.21～ 4.37 mg/m3),季节变化变现为秋季＞冬季＞春季＞夏季.微微型浮游植物对浮游植物总生物量的贡献年平均为5.32％(1.84％～ 8.91％),春季最高,秋季最低.温度在较冷季节(冬春季)里是影响聚球藻蓝细菌生长和分布的控制因素.总之,在近岸环境里,微微型浮游植物并不占优势.     

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.