Responses of diatoms to iron-enrichment (SEEDS) in the western subarctic Pacific, temporal and spatial comparisons

Phytoplankton species composition was analyzed inside and outside of the iron-enriched patch during the SEEDS experiment. Before the iron-enrichment, the phytoplankton community consisted of similar proportions of pico-, nano- and micro-sized phytoplankton, and the micro-phytoplankton was dominated by the pennate diatom Pseudo-nitzschia turgidula. Although all the diatoms, except the nano-sized Fragilariopsis sp., increased during the two weeks of the observation period, the flora in the patch dramatically changed with the increase of phytoplankton biomass to a centric diatom-dominated community. Neritic diatoms, especially Chaetoceros debilis, showed higher growth rates than other diatoms, without any delay in the initiation of growth after the enrichment, and accounted for 90% of the micro-phytoplankton after day 9. In contrast, the oceanic diatoms showed distinct delays in the initiation of growth. We conclude that the responses of the diatoms to the manipulation of iron concentration were different by species, and the fast and intensive response of the phytoplankton to iron-enrichment resulted from the presence of a small amount of neritic diatoms at the study site. The important factors that determine the dominant species in the bloom are the potential growth rates under an iron-replete condition and the growth lag. Abundant species in the patch are widely distributed in the North Pacific and their relative contributions in the Oyashio area and at Stn KNOT are high from spring to summer. However, a characteristic difference of species composition between the SEEDS bloom and natural blooms was the lack of Thalassiosira and Coscinodiscus species in the patch, which usually account for a major part of the phytoplankton community under blooming conditions in the western North Pacific.     

Population dynamics of phytoplankton, heterotrophic bacteria, and viruses during the spring bloom in the western subarctic Pacific

We characterized the community composition of phytoplankton in the western subarctic Pacific from the pre-bloom to the decline phase of the spring bloom with special reference to decreases in the silicic acid concentration in surface waters as an index for diatom bloom development. Furthermore, responses of heterotrophic bacteria and viruses to the spring bloom were also concomitantly investigated. Under pre-bloom conditions when nutrients were abundant but the surface mixed layer depth was relatively deep, chlorophyll (Chl) a concentrations were consistently low and green algae (chlorophytes and prasinophytes), cryptophytes, and diatoms were predominant in the phytoplankton assemblages as estimated by algal pigment signatures. Together with the shallowing of the mixed layer depth and the decrease in silicic acid concentration, diatoms bloomed remarkably in the Oyashio region, though the magnitude of the bloom in the Kuroshio-Oyashio transition (hereafter Transition) region was relatively small. A total of 77 diatom species were identified, with the bloom-forming diatoms mainly consisting of Thalassiosira, Chaetoceros, and Fragilariopsis species. It has become evident that the carotenoid fucoxanthin can serve as a strong indicator of the diatom carbon biomass during the spring diatom bloom. Differences in the species richness of diatoms among stations generally enabled us to separate the Oyashio bloom stations from the Transition and the Oyashio pre-bloom stations. Relatively high values of the Shannon-Wiener index for the diatom species were also maintained during the Oyashio bloom, indicating that a wide variety of species then shared dominance. In the decline phase of the Oyashio bloom when surface nutrient concentrations decreased, senescent diatom cells increased, as inferred from the levels of chlorophyllide a. Although the cell density of heterotrophic bacteria changed little with the development of the diatom bloom, viral abundance increased toward the end of the bloom, suggesting an increased likelihood of mortality among organisms including diatoms resulting from viral infection. This is the first report on the microbial trophodynamics, including viruses, during the spring diatom bloom in the western subarctic Pacific.     

Response of microzooplankton (protists and small copepods) to an iron-induced phytoplankton bloom in the Southern Ocean (EisenEx)

The dynamics, composition and grazing impact of microzooplankton were studied during the in situ iron fertilisation experiment EisenEx in the Antarctic Polar Frontal Zone in austral spring (November 2000). During the 21 day experiment, protozooplankton and small metazooplankton were sampled from the mixed layer inside and outside the patch using Niskin bottles. Aplastidic dinoflagellates increased threefold in abundance and biomass in the first 10 days of the experiment, but decreased thereafter to values twofold higher than pre-fertilisation values. The decline after day 10 is attributed to increasing grazing pressure by copepods. They also constrained ciliate abundances and biomass which were higher inside the fertilised patch than outside but highly variable. Copepod nauplii abundance remained stable whereas biomass doubled. Numbers of copepodites and adults of small copepod species (<1.5 mm) increased threefold inside the patch, but doubled in surrounding waters. Grazing rates estimated using the dilution method suggest that microzooplankton grazing constrained pico- and nanoplankton populations, but species capable of feeding on large diatoms (dinoflagellates and small copepods including possibly nauplii) were selectively predated by the metazoan community. Thus, iron fertilisation of a developing spring phytoplankton assemblage resulted in a trophic cascade which favoured dominance of the bloom by large diatoms.     

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.     

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.     

围隔实验研究台湾海峡2005年夏季小型浮游动物对硅藻水华的摄食

为了研究小型浮游动物对近岸浮游植物藻华的摄食调控作用,于2005年7月,应用"稀释法"并结合高效液相色谱(HPLC)光合色素分析技术,研究了台湾海峡船基围隔实验条件下浮游植物生长率及小型浮游动物摄食率的日变动。结果表明:由于营养盐添加的影响,迅速形成了以尖刺伪菱形藻(Pseudo-nitzschia pungens)为优势种的藻华,生物量(叶绿素a)从实验初始7月6日的1.45μg/dm3迅速增加到7月8日的29.80μg/dm3,随后消退。镜检和光合色素分析的结果显示,实验期间一直以此硅藻占绝对优势。浮游植物的生长率在藻华峰值(7月8日)前保持了较高的生长速率(>1.0/d)且大于小型浮游动物的摄食率;小型浮游动物的摄食率也逐渐增加,7月7日时达到0.86/d,显示有57%以上的浮游植物现存量被摄食。7月8日后,水华迅速消退,摄食率除13日外,均大于浮游植物的生长率。小型浮游动物主要由急游虫(Strombidium spp.)、侠盗虫(Strobilidium spp.)等无壳纤毛虫、异养甲藻-螺旋环沟藻(Gyrodinium spirale)及砂壳纤毛虫等组成,其对浮游植物的生长迅速作出了反应,各类群的丰度在水华峰值后的7月9日均几达最大值,水华后期(11日)大型的无壳纤毛虫达最大值。小型浮游动物的这种组成及变动特点是其保持较高摄食率及一定程度上控制和促进藻华消退的原因之一。     

Mesozooplankton responses to iron-fertilization in the western subarctic Pacific (SEEDS2001)

A mesoscale iron-fertilization experiment was carried out in the western subarctic Pacific during summer 2001. The iron-patch was traced for 14 days after the fertilization, and the abundance and behavior of mesozooplankton were compared with those outside of the patch. The phytoplankton biomass in the patch rapidly increased to over 15 times the initial level by the later half of the observation period, and was composed of large-sized (>10 mm), centric diatoms. Dominant zooplankton species in the upper 200-m depth were large copepods: Neocalanus plumchrus, Neocalanus cristatus, Eucalanus bungii and Metridia pacifica. Mesozoplankton biomass as well as species composition did not change significantly in the patch over the observation period. Furthermore, no changes of vertical distribution or diel vertical migration were observed for any species or stages of mesozooplankton throughout the observation period. However, the abundance of the first copepodite stages of N. plumchrus and E. bungii increased several fold in the patch after the diatom bloom formation compared to the densities outside the patch. The increases of both species are considered to be due to lowered mortality during the egg and nauplius stages. Spawning of N. plumchrus takes place at depth using lipid storage, while spawning of E. bungii takes place in the surface layer supported by grazing. These facts suggest that the relative importance of nauplii in the diets of the large copepods was decreased in the patch by the diatom bloom. Gut-pigment contents of dominant copepods in the patch increased 4–18 times, and the maximum values were observed during the bloom peak. However, the grazing impact on phytoplankton was low throughout the experiment, especially during the bloom period (<6% of the primary production).     

Long-term changes in the seasonality of selected diatoms related to grazers and environmental conditions

The decoupling of trophic interactions could be one of the severe consequences of climate warming in aquatic systems. The timing of phytoplankton blooms, in particular, can affect competition within the plankton community as well as food-web interactions with zooplankton and fish. Using long-term data from Helgoland Roads in the southern North Sea, we examine diatom seasonality, using three representative diatom species combined with environmental and copepod time series over the last four decades. The long-term annual abundances of Guinardia delicatula, Thalassionema nitzschioides and Odontella aurita exhibited interannual variations and dissimilar cyclic patterns during the time period under study (1962-2008). Of the three diatoms, G. delicatula showed a significant trend towards earlier bloom timings for 1962-2008 and a later decline of its abundance over time was found. Grazing and water transparency explained most of the bloom timing fluctuations of the diatoms considered. The annual timing of occurrences of each diatom species was correlated with their preceding concentrations. Earlier bloom timings occurred when autumn/winter concentrations were higher than average and later bloom timings occurred when autumn/winter concentrations were lower than average. Different environmental and predation variables related to the diatom bloom timings were found suggesting that climate warming might not affect the onset of the blooms of the three diatom species in the same manner. The results of the multiple linear regression analyses showed that the timings of decline of the three diatoms were mainly correlated with decreasing nutrient concentrations. Sunshine duration could prolong the duration of the blooms of T. nitzschioides and O. aurita provided that enough nutrients were available. In the case of G. delicatula, however, sunshine duration was negatively correlated with its end of the growth period. G. delicatula and T. nitzschioides showed later decreases in abundances under warmer spring and summer temperatures. Such species specific differences in the sensitivity to the forcing variables could lead to shifts in community structure and could ultimately have wider implications to the overall ecosystem health of the North Sea.     

An in situ iron-enrichment experiment in the western subarctic Pacific (SEEDS): Introduction and summary

To test the iron hypothesis in the subarctic Pacific Ocean, an in situ iron-enrichment experiment (SEEDS) was performed in the western subarctic gyre in July–August 2001. About 350 kg of iron (as acidic iron sulfate) and 0.48 mol of the inert chemical tracer sulfur hexafluoride were introduced into a 10-m deep surface mixed layer over an 80 km² area. This single iron infusion raised dissolved iron levels to 2.9 nM initially. Dissolved iron concentrations rapidly decreased after the infusion, but levels remained close to 0.15 nM even at the end of the 14-day experimental period. During SEEDS there were iron-mediated increases in chlorophyll a concentrations (up to 20 μg l⁻¹), primary production rates, biomass and photosynthetic energy conversion efficiency relative to waters outside the iron-enriched patch. The rapid and very high accumulation of phytoplankton biomass in response to the iron addition appeared to be partly attributable to shallow mixed-layer depth and moderate water temperature in the western subarctic Pacific. However, the main reason was a floristic shift to fast-growing centric diatom Chaetoceros debilis, unlike the previous iron-enrichment experiments in the equatorial Pacific and the Southern Ocean, in both of which iron stimulated the growth of pennate diatoms. The iron-mediated blooming of diatoms resulted in a marked consumption of macronutrients and drawdown of pCO₂. Biological and physiological measurements indicate that phytoplankton growth in the patch became both light- and iron-limited, making phytoplankton biomass relatively constant after day 9. The increase in microzooplankton grazing rate after day 9 also influenced the net growth rate of phytoplankton. There was no significant increase in the export flux of carbon to depth during the 14-day occupation of the experimental site. The export flux between day 4 and day 13 was estimated to be only 13% of the integrated primary production in the iron-enriched patch. The major part of the carbon fixed by the diatom bloom remained in the surface mixed layer as biogenic particulate matter. Our findings support the hypothesis that iron limits phytoplankton growth and biomass in a ‘bottom up’ manner in this area, but the fate of algal carbon remains unknown.     

Meso- and microzooplankton responses to an in situ iron fertilization experiment (SEEDS II) in the northwest subarctic Pacific

A mesoscale iron fertilization experiment was carried out in the western subarctic Pacific during summer 2004. The iron-patch was traced for 26 days after the enrichment, and the abundance and behavior of meso- and microzooplankton was compared with those outside of the patch. The surface chlorophyll-a concentration in the patch was high between days 10 and 13 (2.5 mg m⁻³) and decreased to the initial level after day 20. Microzooplankton grazing rates, estimated by a dilution method, was mostly balanced with phytoplankton growth rates throughout the observed period. Dominant mesozooplankton species in the upper 200 m were copepods: dominated by Eucalanus bungii, Neocalanus plumchrus and Metridia pacifica. Species composition did not change in the patch over the observation period. The copepod biomass was 3–5 times higher than in Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study (SEEDS), the previous iron-enrichment experiment in the same area, before the bloom, and exponentially increased both inside and outside the patch, which was mainly brought by the development of N. plumchrus. The development rates of N. plumchrus were not significantly different between inside and outside the patch. Estimated grazing rate suggest that the copepod grazing was main cause of the low accumulation of phytoplankton biomass, and dominance of grazing-resistant organisms such as large ciliates, large diatoms and diatoms with extremely long setae. “Arrested migration” for M. pacifica and upward shift of vertical distribution by E. bungii were observed during the bloom period, even if the accumulation of phytoplankton biomass was very low compared to other iron-enrichment experiments. These results indicate that the copepod grazing shaped the food-web structure of the lower trophic levels (biomass and species composition) in SEEDS II.     

Nitrogen, Silicon and Zooplankton Controlling the Baltic Spring Bloom: An Experimental Study

The importance of nitrogen, silicon and grazing for the development of the planktonic spring bloom of the transition area between the Baltic Proper and the Gulf of Finland was studied in a laboratory experiment. Water for this experiment was collected prior to the onset of the spring bloom in early April 1991. At this time, inorganic nutrient concentrations were still at their annual maxim. In the experiment, conducted in plastic tanks, the nitrate level was doubled and silicate level tripled by single and combined additions. One tank was filtered with a 100 μm net to exclude large grazers of phytoplankton. Two unmanipulated controls were used. The tanks were maintained at +2 °C for a 3 week period in 12 h of daylight (60 μE m⁻²s⁻¹), during which time their nutrient and phytoplankton dynamics were followed. All added inorganic nitrogen was swiftly taken up inducing rapid phytoplankton growth; consequently, available phosphorus must have met the needs of growth. The results showed the limiting nature of nitrogen on the maximum primary productivity and chlorophyll aconcentration. Silicate additions affected the structure of the phytoplankton assemblage by promoting specifically the growth of the diatomChaetoceros holsaticusSchütt, which produced resting spores at later stages of the bloom, whileChaetoceros wighamiiBrightwell was the dominant diatom species in other tanks. The absence of large grazers had no effect on phytoplankton dynamics.     

Enhanced carbohydrate production by Southern Ocean phytoplankton in response to in situ iron fertilization

Storage carbohydrates (e.g., water-extractable β-1,3-d-glucan in diatoms) are of key importance for phytoplankton growth in a variable light climate, because they facilitate continued growth of the cells in darkness by providing energy and carbon skeletons for protein synthesis. Here, we tested the hypothesis that synthesis of storage carbohydrates by phytoplankton in the Southern Ocean is reduced by low iron and light availability. During the EisenEx/CARbon dioxide Uptake by the Southern Ocean (CARUSO) in situ iron enrichment experiment in the Atlantic sector of the Southern Ocean in November 2000, we studied the dynamics of water-extractable carbohydrates in the particulate fraction over the period of 3 weeks following the iron release. The areal amount (integral between 0- and 100-m depth) of carbohydrates increased from 1400 to 2300 mg m⁻² inside the iron-enriched patch, while remaining roughly constant in the surrounding waters. Most of the increase inside the patch was associated with the fraction of large (>10 μm) phytoplankton cells, consistent with the shift in the community structure towards larger diatoms. Deck incubations at 60% of the ambient irradiance revealed that the diurnal chlorophyll a (Chl a)-specific production rates of water-extractable polysaccharides were significantly higher for “in-patch” than for “out-patch” samples (0.5 vs. 0.3 μg C [μg Chl a]⁻¹ h⁻¹, respectively). Together with the higher photochemical efficiency of photosystem II (F_v/F_m), this indicates enhanced photosynthetic performance in response to iron fertilization. In addition, the nocturnal polysaccharide consumption rates were also enhanced by iron release, causing a striking increase in the diel dynamics of polysaccharide concentration. An iron-stimulated increase in diel dynamics was also observed in the fluorescence and size of pico- and nanophytoplankton cells (measured by flow cytometry) and is indicative of enhanced phytoplankton growth. Diurnal polysaccharide production by phytoplankton inside the patch was light-limited when they were incubated at intensities below ca. 200 μmol m⁻² s⁻¹ (daytime average). These irradiance levels correspond to those at 20- to 30-m depth in situ, whereas the upper mixed layer was frequently several-fold deeper due to storms. Therefore, these first measurements of phytoplankton carbohydrates during an in situ iron release experiment have revealed that both light and iron availability are the key factors controlling the synthesis of storage carbohydrates in phytoplankton and, hence, the development of diatom blooms in the Southern Ocean.     

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.     

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.     

Biological controls on dissolution of diatom frustules during their descent to the deep ocean: Lessons learned from controlled laboratory experiments

The majority of opal produced by diatoms dissolves during their sedimentation to the seafloor, but spatial and temporal variability of dissolution rates are large. Controlled laboratory experiments using live phytoplankton or phyto-detritus may help identify the different processes, including those that are biologically mediated or physico-chemically driven, that impact the dissolution of frustules and the aforementioned variability. Results of eight bSiO₂ dissolution experiments, seven of which were conducted at low temperatures (<6 °C) are presented within the context of earlier similar studies, and different phases of dissolution dynamics characterized. TEP concentration, aggregation and the physiological status of the diatoms determined the period during which diatoms may maintain the protective membrane that surrounds their frustule and effectively reduces or completely inhibits (lag period) dissolution for some time. Once diatoms loose the capability to maintain their protective membrane, bacterial activity compromises it. Physico-chemical dissolution, which depends on frustule structure and abiotic environmental conditions, begins once the protective membrane is damaged. The ability of diatoms to maintain their membrane, the bacterial composition and activity governing its degradation, and the physico-chemical dissolution dynamics of exposed frustules are all impacted by temperature. In our experiments instantaneous dissolution rates were not dependant on bSiO₂ concentration at low temperatures, although such a relationship was observed under otherwise identical conditions at 15 °C, implying that biotic factors rather than physico-chemical processes initially dominated dissolution at polar temperatures. Since inhibition of bSiO₂ dissolution at low temperatures was inhibited to a greater extent than organic matter degradation, we postulate that it was not reduced bacterial activity but the enhanced ability of diatoms to maintain their membrane and thus withstand microbial attack that caused the low initial dissolution rates at <6 °C. In situ, interactions between the different biotic and abiotic processes impacting dissolution combined with differences in sinking velocity of diatom aggregates and grazing effects could easily explain high spatial and temporal variability in the accumulation of diatoms on the seafloor. Simple calculations based on our experimental results suggest that Fragilariopsis kerguelensis, for example, would be appreciably more likely to reach the seafloor than Chaetoceros debilis if both grow at low growth rates, e.g. under growth limiting conditions. However, dissolution behavior of Chaetoceros debilis during sedimentation may differ under conditions where this species forms large blooms.     

A daily study of the diatom spring bloom at Roscoff (France) in 1985. I. The spring bloom within the annual cycle

The repeated occurrence of a monospecific bloom of the plankton diatom Rhizosolenia delicatula at Roscoff (western English Channel) was made the subject of an interdisciplinary research programme. Samples were taken at daily intervals from April to July and at longer intervals during the remaining part of the year 1985. Routine physical parameters, light transmission, nutrients, dissolved oxygen, particle load, particulate N and P, chlorophyll content, phytoplankton counting, and zooplankton biomass were measured as a basis for more specific studies (to follow as subsequent papers in this series).The area is characterized by: high tidal range, permanent mixing throughout the year, low attenuation coefficients, moderate nutrient supply, and the dominance of benthic algae over phytoplankton. The spring bloom is significantly delayed with respect to the usual model for the temperate seas. Tidal cycles are expected to exert the main influence on bloom dynamics at the time scale of phytoplankton growth.     

Role of dissolved silicate in the occurrence of a phytoplankton bloom

The spring bloom of phytoplankton was studied in March in Funka Bay, Japan, to test the Tsunogai (1979)'s hypothesis regarding the role of silicate in the bloom. The hypothesis comprises two parts. 1) Diatoms are predominant when all the physical and chemical conditions are adequate for plankton growth. 2) Since the Si:P ratio of the diatom body is usually much larger than that of sea water, flagellates (non-siliceous phytoplankton) replace diatoms after dissolved silicate in the sea water has been almost completely consumed by diatoms. At the end of the bloom in late March phosphate still remained in the water but silicate was exhausted and the main species of phytoplankton changed from diatoms to flagellates. Grazing pressure by zooplankton at this time was not so great. A model using the data on assimilation rates of silicate showed a dramatic change of silicate uptake in late March. Poison in scallops caused byProtogonyaulux sp. (dinoflagellates) rapidly increased from mid-April at all stations along the coast of Funka Bay. All of these findings support Tsunogai's hypothesis.     

Factors controlling the temporal variation of fatty acids in piculate matter during a phytoplankton bloom in a marine mesocosm

Fatty acids present in suspended particles were examined following the onset of a phytoplankton bloom in an experimental marine mesocosm set up in Patricia Bay, Saanich Inlet, British Columbia, Canada. The predominantly diatom bloom, triggered by the addition of nutrients at the commencement of the experiment, was succeeded by several other phytoplankton forms, which in turn led to changes in particulate chl a, and organic carbon, and nitrogen within the experimental enclosure. An examination of individual fatty acids indicated that the variation of individual fatty acids can be classified into three groups, although the variation pattern of individual fatty acids differed from one another. An attempt was made to establish the factors controlling particulate fatty acid concentrations by principal component analysis. After Varimax rotation, six factors were extracted, of which four factors contributed to 83% of the data matrix. These were mainly dependent on the source of the particles i.e., diatoms, dinoflagellates, and bacteria.Fatty acid concentrations obtained by analysis of different lipid class fractions indicated that the production of fatty acids was distinctly related to the growth stage of the phytoplankton bloom. Fatty acids in triglycerides peaked during stationary phase, whereas fatty acids in polar lipids were high during the stage of an increase of phytoplankton biomass. However, fatty acid composition in both lipid classes changed under the influence of species succession of the phytoplankton population. This strongly suggested that species succession of the phytoplankton exerted the most significant control on fatty acid composition of particulate matter.     

Growth and succession patterns of major phylogenetic groups of marine bacteria during a mesocosm diatom bloom

Our objective was to track microbial processes associated with serial degradation of organic matter derived from algal blooms. To do this, we analyzed population fluctuations and growth responses of major phylogenetic groups of free-living marine bacteria. We used bromodeoxyuridine immunocytochemistry–fluorescence in situ hybridization methodology to examine marine bacterial community development during and after a diatom bloom in a mesocosm. We revealed that the Roseobacter/Rhodobacter, SAR11, Alteromonas, and Bacteroidetes groups were clearly major phylotypes responsible for most free-living bacterial biomass and production throughout the experiment. The clearest bacterial response was a proliferation of the Alteromonas group (cells with large volumes) during development of the bloom (up to 30?% of actively growing cells). Populations of these bacteria declined sharply thereafter, likely due to grazing. Alteromonas group responses suggest that these bacteria strongly influenced the flux of organic matter at an early bloom stage. The growth potential of Bacteroidetes was relatively large as the bloom peaked; this early development probably contributed to the initial stage of bloom decomposition. In contrast, the contribution of Roseobacter/Rhodobacter to total bacterial production increased at a late stage of decomposing of the bloom. The contributions of Betaproteobacteria, SAR11, and SAR86 groups to total bacterial abundance and production were relatively minor throughout the experiment. These results imply that the ability to utilize organic matter derived from diatoms varies among bacterial phylotypes, and, frequently, less abundant but ecological specialist taxa such as Alteromonas may play major roles in the flux of organic matter during diatom blooms.     

围隔生态系内不同粒级浮游植物群落变化研究

通过现场围隔实验,模拟赤潮发生过程,研究了种群不同生长阶段中不同粒级浮游植物种群的变化情况。结果表明,添加营养盐能有效促进浮游植物的生长,东海原甲藻围隔（M1）和自然水体围隔（M2）中浮游植物分别于第7天和第4天出现生长高峰,叶绿素a最大值分别为112．79mg／m和235．60mg/m。微型浮游植物与微微型浮游植物存在竞争,微型对微微型生长的抑制作用：M2〉M1。在营养盐丰富时,硅藻的增殖速率比东海原甲藻快,达到高峰期时间短,消亡也快。硅的减少促进硅藻水华的消亡。