Fatty acids associated with the frustules of diatoms and their fate during degradation—A case study in Thalassiosira weissflogii

Diatoms are major actors in the export of organic carbon out of the euphotic zone. Yet, the processes linking biogenic silica and carbon sedimentation fluxes to deep oceanic layers remain unclear. Analysing organic fractions in biominerals is challenging because efficient cleaning often led to structural alteration of organic molecules. Hence, although lipids are widely used as biogeochemical markers in ocean flux study, few studies have dealt with the lipids that are associated with frustules. In the present study, a protocol was set up to extract and quantify the fatty acids associated to the frustule of the diatom species Thalassiosira weissflogii. The protocol involves solvent extraction of diatom external lipids, followed by clean frustule dissolution by 4% NaOH during 1 h at 95 °C and subsequent solvent re-extraction of frustule-associated lipids. Results confirmed that this protocol was efficient first, to isolate the frustule from the rest of the cellular organic carbon and second to extract and quantify fatty acids (FA) associated to frustules of this species. FA composition of the frustules was significantly different from that of the whole cells consisting primarily of 14:0, 16:0 and 18:0 FA, as well as a smaller portion of 16:1 and 18:1 unsaturated FA. Frustule-associated FA constituted 7% of the total FA and 1.8% of the total POC. The 30 days T. weissflogii degradation/dissolution experiment suggested that frustule FA 14:0 and 16:0 were mainly associated with the bSiO₂ phase dissolving slowly as no degradation of this pool was measured despite 78% frustule dissolution. At the end of the degradation experiment, this pool constituted 5.8% of the remaining total POC suggesting an effective protection by the frustule through strong interaction with the biogenic silica which is consistent with the correlation observed at depth between Si and POC sedimentation fluxes.     

Mechanisms determining species dominance in a phytoplankton bloom induced by the iron fertilization experiment EisenEx in the Southern Ocean

The dynamics of phytoplankton species populations recorded during the 3-week, iron-fertilization experiment EisenEx carried out in spring in the Antarctic Polar Frontal Zone are presented and discussed as the difference between growth and mortality rates. Only two cosmopolitan diatom species, the centric Chaetoceros debilis and the pennate Pseudo-nitzschia lineola, increased population density exponentially throughout the experiment to 150- and 90-fold of initial values, respectively. Because C. debilis initial abundance was tenfold lower than that of P. lineola, the two contributed 1% and 21% to bloom biomass, respectively at the end of the experiment, high-lighting the role of seeding in bloom formation. The other significant species increased population size at a linear rate throughout the experiment or for a short spurt phase to 3- to 18-fold of initial values. Conservative estimates of mortality rates within diatom species populations were obtained by comparing net accumulation rates of full cells with those of empty and broken frustules. The ratios were consistent over time for the various species but varied widely between them. The species-specific variation can be explained by differences in both growth and mortality rates, the latter partly due to either selective grazing or avoidance by the large protozoo- and metazooplankton populations present. Selective predation by the abundant copepod populations on protistan grazers of diatoms (ciliates and heterotrophic dinoflagellates) apparently aided diatom biomass build-up. The response patterns of populations of the phytoplankton species present fall into six categories comprising disparate species, indicating that phylogeny is a poor predictor of ecology. The group that did not respond to fertilization was the most diverse and included both endemic and cosmopolitan as well as background and bloom-forming species. This lack of response to the advent of favorable growth conditions indicates that proximate factors during EisenEx triggered growth only in some species but had little effect on others. We attribute the differences in behavior to ultimate factors such as seasonal effects on life cycles and other internal constraints on growth rates. The implications for our understanding of the evolutionary ecology of phytoplankton and its impact on global biogeochemical cycles are pointed out.     

The use of soft X-ray spectromicroscopy to investigate the distribution and composition of organic matter in a diatom frustule and a biomimetic analog

Diatoms play a significant role in the global carbon cycle through their role in biogenic silica production and the transport of organic matter to the seafloor. Recent work has shown that silicified diatom frustules contain a significant amount of organic matter, and that the proportion of diatom-bound organic matter increases with depth in the water column and sediments. Here, we investigate the association between organic matter and the mineral phase. We used a combination of scanning transmission X-ray microscopy (STXM) and carbon X-ray absorption near-edge structure (XANES) spectroscopy to characterize the distribution and composition of organic matter in frustules of the diatom Cylindrotheca closterium and a biomimetic silica gel. To our knowledge, this study represents the first successful attempt to simultaneously image and obtain chemical information about the organic matter within a diatom frustule using X-ray spectromicroscopy near the carbon edge. Organic carbon, most likely protein, was distributed throughout the frustules and was not removed by harsh chemical treatment. The physical structure of the frustules appeared to be related to the chemical composition of this organic matter, with aromatic or unsaturated carbon being concentrated in the most intricately patterned regions of the frustule. A similar physical and chemical structure was observed in a biomimetic silica gel precipitated spontaneously with polylysine. These results are consistent with the theory that organic constituents of diatom frustules direct silica precipitation and become incorporated within the silica matrix as it forms. The relationship between organic matter composition and silica morphology, the failure of harsh chemical treatments to remove this organic matter, and the spontaneous nature of the co-precipitation of silica and organic matter indicate some chemical interaction between the siliceous and organic components of diatom frustules. Frustule-bound organic matter should therefore be protected from decomposition in the water column or diagenetic alteration in sediments unless the frustule dissolves.     

Settling fluxes of diatoms to the interior of the Antarctic circumpolar current along 170 °W

An array of four sediment trap moorings recorded the particulate flux across the Antarctic Circumpolar Current (ACC) at 170 °W, between November 1996 and January 1998, as part of the US JGOFS-Antarctic Environment and Southern Ocean Process Study (AESOPS) program. The trap locations represent sampling within the Polar Frontal Zone, the Antarctic Polar Front, the Antarctic Zone and the Southern Antarctic Zone. Here we report observations from 1000 m below the sea-surface compared to seafloor and surface water distributions. Sub-sample splits from each trap were obtained and total diatom flux and species composition were determined. The diatom fluxes were quantified using both a dilution and a ‘spike’ method to allow for the rapid repeatability of measurements. Diatom flux was found to be highly seasonal across the ACC particularly at higher latitudes. Marine snow aggregates of intact diatom cells and chains were the major components of the biogenic flux. Siliceous particle size was noted to decrease with increasing latitude, which could be aligned with a shift of the diatom assemblage to small-size species/sea-ice affiliated species. A ‘double-structured’ diatom flux was recorded at the location of the Antarctic Polar Front trap, with a shift in the diatom assemblage from larger to smaller diatoms in the second flux episode. The sediment trap assemblage shows deviations from the surface water assemblage, while surface sediment samples indicate that significant dissolution occurs after 1000 m and at the sediment–water interface. Estimation of diatom biovolumes across the ACC shows that large diatoms have the potential to greatly impact biogenic fluxes to the ocean interior despite their low fluxes. Small species of the genus Fragilariopsis could potentially export as much C_org as Fragilariopsis kerguelensis near the retreating ice edge. However, their low abundance in the surface sediments also suggests that these diatoms are a shallow export species.     

2014年春季季风间期东印度洋赤道及其邻近海域硅藻群落

2014年4月10日至5月13日在东印度洋赤道区及其邻近海域(10.08°N-6.00°S,80.00°~96.10°E)进行硅藻物种组成和群落结构的调查。分析了45个网采样品,共鉴定出浮游硅藻34属113种(包括变种、变形及未定名种),大部分物种为热带外洋性种以及暖海外洋性种,与该海区的热带及亚热带的环境特征一致。优势种为佛氏梯形藻(Climacodium frauenfeldianum)、地中海细柱藻(Leptocylindrus mediterraneus)、密聚角毛藻(Chaetoceros coarctatus)、美丽漂流藻(Planktoniella foromsa)、大西洋角毛藻那不勒斯变种(Chaetoceros atlanticus var. neapolitanus)、距端假管藻(Pseudosolenia calcar-avis,即距端根管藻Rhizosolenia calcaravis)、圆柱几内亚藻(Guinardia cylindrus)、达蒂角毛藻(Chaetoceros dadayi)、伏氏海线藻(Thalassionema frauenfeldii)、离心列海链藻(Thalassiosira excentrica)、瘤面角毛藻(Chaetoceros bacteriastroides)以及笔尖根管藻粗径变种(Rhizosolenia styliformis var. latissima)等。硅藻的平均细胞丰度为1.855×103个/m3,其平面分布不均匀,赤道断面细胞丰度较低,高值区出现在海区北部。聚类分析发现有7种生态类群,这些硅藻的种类和丰度平面分布与上层水体(200 m以浅)温度、盐度及营养盐的水柱平均值有较好的对应关系。     

Biogeochemical responses to late-winter storms in the Sargasso Sea,III—Estimates of export production using 234Th:238U disequilibria and sediment traps

Direct measurements of new production and carbon export in the subtropical North Atlantic Ocean appear to be too low when compared to geochemical-based estimates. It has been hypothesized that episodic inputs of new nutrients into surface water via the passage of mesoscale eddies or winter storms may resolve at least some of this discrepancy. Here, we investigated particulate organic carbon (POC), particulate organic nitrogen (PON), and biogenic silica (BSiO₂) export using a combination of water column ²³⁴Th:²³⁸U disequilibria and free-floating sediment traps during and immediately following two weather systems encountered in February and March 2004. While these storms resulted in a 2–4-fold increase in mixed layer NO₃ inventories, total chlorophyll a and an increase in diatom biomass, the systems were dominated by generally low ²³⁴Th:²³⁸U disequilibria, suggesting limited particle export. Several ²³⁴Th models were tested, with only those including non-steady state and vertical upwelling processes able to describe the observed ²³⁴Th activities. Although upwelling velocities were not measured directly in this study, the ²³⁴Th model suggests reasonable rates of 2.2–3.7 m d^?1.Given the uncertainties associated with ²³⁴Th derived particle export rates and sediment traps, both were used to provide a range in sinking particle fluxes from the upper ocean during the study. ²³⁴Th particle fluxes were determined applying the more commonly used steady state, one-dimensional model with element/²³⁴Th ratios measured in sediment traps. Export fluxes at 200 m ranged from 1.91±0.20 to 4.92±1.22 mmol C m^?2 d^?1, 0.25±0.08 to 0.54±0.09 mmol N m^?2 d^?1, and 0.22±0.04 to 0.50±0.06 mmol Si m^?2 d^?1. POC export efficiencies (Primary Production/Export) were not significantly different from the annual average or from time periods without storms, although absolute POC fluxes were elevated by 1–11%. This increase was not sufficient, however, to resolve the discrepancy between our observations and geochemical-based estimates of particle export. Comparison of PON export rates with simultaneous measurements of NO₃^? uptake derived new production rates suggest that only a fraction, <35%, of new production was exported as particles to deep waters during these events. Measured bSiO₂ export rates were more than a factor of two higher (p<0.01) than the annual average, with storm events contributing as much as 50% of annual bSiO₂ export in the Sargasso Sea. Furthermore it appears that 65–95% (average 86±14%) of the total POC export measured in this study was due to diatoms.Combined these results suggest that winter storms do not significantly increase POC and PON export to depth. Rather, these storms may play a role in the export of bSiO₂ to deep waters. Given the slower remineralization rates of bSiO₂ relative to POC and PON, this transport may, over time, slowly decrease water column silicate inventories, and further drive the Sargasso Sea towards increasing silica limitation. These storm events may further affect the quality of the POC and PON exported, given the large association of this material with diatoms during these periods.     

Particle fluxes in San Pedro Basin, California: A four-year record of sedimentation and physical forcing

Moored sediment traps were deployed from January 2004 through December 2007 at depths of 550 and 800 m in San Pedro Basin (SPB), CA (33°33.0′N, 118°26.5′W). Additionally, floating sediment traps were deployed at 100 and 200 m for periods of 12-24 h during spring 2005, fall 2007, and spring 2008. Average annual fluxes of mass, particulate organic carbon (POC), ??¹³C_org, particulate organic nitrogen (PON), ??¹⁵N-PON, biogenic silica (bSiO₂), calcium carbonate (CaCO₃), and detrital material (non-biogenic) were coupled with climate records and used to examine sedimentation patterns, vertical flux variability, and organic matter sources to this coastal region. Annual average flux values were determined by binning data by month and averaging the monthly averages. The average annual fluxes to 550 m were 516±42 mg/m² d for mass (sdom of the monthly averages, n=117), 3.18±0.26 mmol C/m² d for POC (n=111), 0.70±0.05 mmol/m² d for CaCO₃ (n=110), 1.31±0.21 mmol/m² d for bSiO₂ (n=115), and 0.35±0.03 mmol/m² d for PON (n=111). Fluxes to 800 and to 550 m were similar, within 10%. Annual average values of ??¹³C_org at 550 m were −21.8±0.2‰ (n=108), and ??¹⁵N averages were 8.9±0.2‰ (n=95). The timing of both high and low flux particle collection was synchronous between the two traps. Given the frequency of trap cup rotation (4-11 days), this argues for particle settling rates ≥83 m/d for both high and low flux periods. The moored traps were deployed over one of the wettest (2004-2005, 74.6 cm rainfall) and driest (2006-2007, 6.6 cm) rain years on record. There was poor correlation (Pearson's correlation coefficient, 95% confidence interval) of detrital mass flux with: C_org/N ratio (r=0.10, p=0.16); ??¹⁵N (r=−0.19, p=0.02); and rainfall (r=0.5, p=0.43), suggesting that runoff does not immediately cause increases in particle fluxes 15 km offshore. ??¹³C_org values suggest that most POC falling to the basin floor is marine derived. Coherence between satellite-derived chlorophyll a records from the trap location (±9 km² resolution) and SST data indicates that productivity and export occurs within a few days of upwelling and both of these parameters are reasonable predictors of POC export, with a time lag of a few days to 2 weeks (with no time lag—SeaWiFS chlorophyll a and POC flux, r=0.25, p=0.0014; chlorophyll a and bSiO₂ flux, r=0.28, p=0.0002).     

Dissolution of silica accompanied by oxygen consumption in the bottom layer of Japan’s central Seto Inland Sea in summer

Distribution of silicic acid (Si(OH)₄) in bottom water was investigated in the central Seto Inland Sea under stratified conditions in summer. Water samples were collected at 10 stations on April 24 and 25 and July 7 and 8, 2012. In July, stratification progressed, and a cold water mass (dome) of <20 °C appeared. In response to the cold dome, low oxygen content was observed in the bottom layer of the eastern part of Hiuchi-Nada. In this water mass site, apparent oxygen utilization values calculated from dissolved oxygen (DO) concentrations increased, coinciding with increase of Si(OH)₄ concentrations from April to July. This suggests that increase of Si(OH)₄ [dissolution of biogenic silica (diatom frustules)] was accompanied by DO consumption due to degradation of organic matter such as plankton soft tissue. These findings suggest that a bacterial degradation of the organic matrix that covers diatom frustules could accelerate the dissolution of biogenic silica in bottom water under stratified conditions.     

Isotopic constraints on the Si-biogeochemical cycle of the Antarctic Zone in the Kerguelen area (KEOPS)

Estimation of the silicon (Si) mass balance in the ocean from direct measurements (Si uptake-dissolution rates …) is plagued by the strong temporal and spatial variability of the surface ocean as well as methodological artifacts. Tracers with different sensitivities toward physical and biological processes would be of great complementary use. Silicon isotopic composition is a promising proxy to improve constraints on the Si-biogeochemical cycle, since it integrates over longer timescales in comparison with direct measurements and since the isotopic balance allows to resolve the processes involved, i.e. uptake, dissolution, mixing. Si-isotopic signatures of seawater Si(OH)₄ and biogenic silica (bSiO₂) were investigated in late summer 2005 during the KEOPS experiment, focusing on two contrasting biogeochemical areas in the Antarctic Zone: a natural iron-fertilized area above the Kerguelen Plateau (< 500 m water depth) and the High Nutrient Low Chlorophyll area (HNLC) east of the plateau (> 1000 m water depth). For the HNLC area the Si-isotopic constraint identified Upper Circumpolar Deep Water as being the ultimate Si-source. The latter supplies summer mixed layer with 4.0 ± 0.7 mol Si m^? 2 yr^? 1. This supply must be equivalent to the net annual bSiO₂ production and exceeds the seasonal depletion as estimated from a simple mixed layer mass balance (2.5 ± 0.2 mol Si m^? 2 yr^? 1). This discrepancy reveals that some 1.5 ± 0.7 mol Si m^? 2 yr^? 1 must be supplied to the mixed layer during the stratification period. For the fertilized plateau bloom area, a low apparent mixed layer isotopic fractionation value (?³⁰Si) probably reflects (1) a significant impact of bSiO₂ dissolution, enriching the bSiO₂ pool in heavy isotope; and/or (2) a high Si uptake over supply ratio in mixed layer at the beginning of the bloom, following an initial closed system operating mode, which, however, becomes supplied toward the end of the bloom (low Si uptake over supply ratio) with isotopically light Si(OH)₄ from below when the surface Si(OH)₄ pool is significantly depleted. We estimated a net integrated bSiO₂ production of 10.5 ± 1.4 mol Si m^? 2 yr^? 1 in the AASW above the plateau, which includes a significant contribution of bSiO₂ production below the euphotic layer. However, advection which could be significant for this area has not been taken into account in the latter estimation based on a 1D approach of the plateau system. Finally, combining the KEOPS Si-isotopic data with those from previous studies, we refined the average Si-isotopic fractionation factor to ? 1.2 ± 0.2‰ for the Antarctic Circumpolar Current.     

Phytoplankton blooms in the Urdaibai estuary during summer: Physico-chemical conditions and taxa involved

In the Urdaibai estuary, despite its small volume in relation to the tidal prism, phytoplankton grows massively in the upper and intermediate zones of the estuary during summer when rainfall is low to moderate. Data obtained in 23 sampling transects along the longitudinal axis of the estuary, undertaken in July 1993 and August 1994, showed three distinct zones in terms of the phytoplankton species composition: in the upper zone the phytoplankton assemblage was dominated by the diatoms Cyclotella spp., the dinoflagellate Glenodinium foliaceum, cryptophytes and euglenophytes; in the intermediate zone the diatom Chaetoceros ceratosporus, the dinoflagellates Peridinium quinquecorne and Prorocentrum minimum and cryptophytes were the most abundant; in the lower zone diatoms such as Leptocylindricus danicus and Skeletonema costatum together with the dinoflagellates Prorocentrum spp. were dominant. The distribution and abundance of these phytoplankton forms is discussed in relation to variations in salinity, water column stratification, nutrients and temperature, which in turn vary mainly as a function of meteorological conditions (i.e. rainfall, irradiance) and tidal amplitude. The longitudinal distribution of phytoplankton cells suggests that the diatoms Cyclotella spp. and Chaetoceros ceratosporus undergo a greater dispersion along the estuary during rain events than the flagellates, thus contributing to the transport of silica to the outer areas of the estuary.     

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.     

Particulate Flux and Cd/P Ratio of Particulate Material in the Pacific Ocean

Time-series Mark 7 sediment traps were deployed at three stations at 0°N, 13°N and 48°N along 175°E to investigate seasonal and spatial variations of particulate material flux. Chemical analysis of particulate material was performed for four major chemical components, viz. opal, CaCO₃, organic material and clay minerals, Cd and P in the particulate material were also determined. We discuss the characteristics of particulate material at each site and the transportation of Cd and P to deep water by the particulate material. The total mass fluxes and variation of fluxes at each site reflect oceanographic conditions, such as biological productivity and kind of major planktonic organisms. At the northern site, large mass fluxes with a spring bloom and high ratios of opal are characteristic. Relatively small mass fluxes with high ratios of CaCO₃ are distinct, and dissolution of CaCO₃ due to sinking is recognized in the middle latitude and 0°N sites. The larger flux at the lower trap than the upper trap at the equatorial site suggests influence by lateral transport in the deep water. Distinctive decreasing Cd/P ratio and CaCO₃ concentrations in the particulate material with increasing depth suggests that the change of Cd/P ratio in the intermediate and deep water occurs through the dissolution of CaCO₃. The dissolved Cd/P ratios in the deep water are proportional to the age of the deep water in the Atlantic but not in the Pacific. This is explained by the difference of kinds of particulate material transporting Cd and P in the deep water between the oceans. That is, the major planktonic organisms are planktons of CaCO₃ tests in the Atlantic Ocean and diatoms of opal tests in the North Pacific Ocean.     

硅藻硅酸盐转运子(silicate transporter, SIT)研究进展

硅藻是海洋浮游植物主要类群和海洋生态系统的重要初级生产者,其硅质壁的存在使其成为了海洋生物硅的最大贡献者,而且成为海洋碳汇的主要来源。因此,硅质壁形成机制的阐明至关重要。硅酸盐转运是硅藻硅质壁形成中的关键,阐明其转运机制对建立硅藻代谢途径模型和进一步理解硅藻在海洋生态系统及生物地球化学循环中的作用具有重大意义。硅酸盐转运子(silicate transporter,SIT)是硅藻中参与硅吸收或转运的一类膜蛋白,因其重要性及多样性成为讨论及研究的焦点。本文综述了硅藻中SIT的分子特征、分子机制、功能多样性及进化关系,为SIT转运机制的研究提供了理论基础。     

Strong CO2 outgassing from high nutrient low chlorophyll coastal waters off central Chile (30°S): The role of dissolved iron

Carbonate system parameters (pH and alkalinity) were used to estimate the coastal water CO₂ fluxes off central Chile (30°S) during September 2007. Coastal waters rich in nitrate and silicate were strongly CO₂ supersaturated and normally poor in chlorophyll a. MODIS satellite chlorophyll a data suggest that phytoplankton biomass remained particularly low during September 2007 although coastal waters were highly fertilized with nitrate and silicate. The phytoplankton gross primary productivity in macronutrient-rich waters was very low with the exception of shallow waters (e.g. within or near bays). Several iron-enrichment bottle experiments show that fCO₂ rapidly decreases during iron-enrichment treatments compared to controls. This suggests that iron limitation of phytoplankton growth (mainly diatoms) plays a role in maintaining high-CO₂ outgassing by preventing rapid interception of upwelled CO₂.     

3种海洋硅藻不同培养阶段挥发性成分的比较分析

本文应用顶空固相微萃取气相色谱-质谱联用技术,对威氏海链藻Thalassiosira weissflogii、小新月菱形藻Nitzschia closterium、角毛藻Chaetoceros calcitrons 3种硅藻的挥发性成分进行分析。结果表明,在3种硅藻的培养过程中,其挥发性成分除了壬醛及醇类呈现先增后降的趋势外,大多数不饱和醛类的含量在不断增加,烷烯烃类物质在不断减少,而乙酸乙酯、2-甲基-2-丙酸甲酯这类酯类化合物在角毛藻衰败期会显著富集。通过对代表性饵料硅藻的挥发性成分进行研究,可为海水养殖过程中微藻投喂种类、投喂时机的选择提供理论依据,并为海洋硅藻的进一步开发奠定基础。     

Interactive effects of iron,irradiance and CO2 on Ross Sea phytoplankton

We conducted a factorial shipboard continuous culture experiment to examine the interactive effects of altered iron, irradiance and CO₂ on the summer phytoplankton community of the Ross Sea, Antarctica. After 18 days of continuous incubation, iron enrichment increased phytoplankton biomass, nutrient drawdown, diatom and Phaeocystis abundance, and some photosynthetic parameters. High irradiance significantly increased the number of Phaeocystis antarctica colonies, as well as P. antarctica abundance relative to diatoms. Iron and light had significant interactive effects on diatom and P. antarctica pigment concentrations, P. antarctica colony abundance, and Si:N, Si:C, and N:P ratios. The major influence of high CO₂ was on diatom community structure, by favoring the large centric diatom Chaetoceros lineola over the small pennate species Cylindrotheca closterium. The ratio of centric to pennate diatoms was significantly responsive to changes in all three variables individually, and to all of their possible two- and three-way combinations. These results suggest that shifts in light, iron, and CO₂ and their mutual interactions all play a role in controlling present day Ross Sea plankton community structure, and need to be considered when predicting the possible future responses of biology and biogeochemistry in this region.     

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.     

2006年冬季北黄海网采浮游植物群落结构

根据2006年12月30日—至2007年1月17日北黄海的调查资料,对该海域浮游植物的种类组成、优势种、丰度及其分布和多样性等基本状况进行了分析。本次调查共鉴定浮游植物4门68属131种,主要以温带近岸和广布性种为主,其中硅藻有53属113种,占总种数的86.3%,甲藻有11属16种,占总种数的12.2%。浮游植物丰度平均值为88.89×104个/m3,硅藻丰度平均值为86.58×104个/m3,甲藻丰度平均值为2.28×104个/m3,硅藻丰度分布趋势决定了浮游植物丰度的分布趋势。辽宁南岸是浮游植物密集区,山东半岛北岸其次,而北黄海中部是浮游植物的稀疏区。优势种为:短角弯角藻(Eucampia zodiacus)、具槽帕拉藻(Paralia sulcata)、尖刺拟菱形藻(Pseu-do-nitzschia pungens)、密连角毛藻(Chaetoceros densus)、柔弱角毛藻(Chaetoceros debilis)、刚毛根管藻(Rhizosolenia setigera)。浮游植物群落Shannon-Weiner物种多样性指数平均值为1.80,Peilou均匀度指数平均值为0.42。与1959年1月相比,2007年1月北黄海浮游植物丰度由150.00×104个/m3降为88.89×104个/m3,下降了近41%,硅藻丰度由148.00×104个/m3降为86.58×104个/m3,而甲藻丰度由1.25×104个/m3上升为2.28×104个/m3,占浮游植物丰度的比例由0.8%上升为2.5%。主要优势种及优势属也发生了一定程度的变化,但浮游植物群落结构仍以硅藻为主、甲藻其次,浮游植物丰度总的分布格局变化不明显。     

2014年夏季南海北部浮游植物群落结构

2014年8月20日-9月6日对南海北部(18°~22°N,114°~116°E)的11个站位进行了浮游植物群落结构的调查,其中2个站位为时间连续站。共鉴定出浮游植物4门68属229种(包括变种、变型及未定种),其中硅藻门43属147种,甲藻门20属75种。浮游植物平均细胞丰度为16.318×103 cells/L,硅藻的平均细胞丰度为14.653×103 cells/L,占总丰度的89.80%,硅藻的分布决定了浮游植物的分布。近岸冲淡水含有丰富的营养盐,因此近岸海域的细胞丰度高于大洋海域,垂直分布表现出先增加后降低的趋势。本次调查的优势种为铁氏束毛藻(Trichodesmium thiebautii)、中肋骨条藻(Skeletonema costatum)、拟脆杆藻(Fragilariopsis sp.)、短孢角毛藻(Chaetoceros brevis)、菱形海线藻(Thalassionema nitzschioides)、柔弱伪菱形藻(Pseudo-nitzschia delicatissima)、尖刺伪菱形藻(Pseudo-nitzschia pungens)、扁面角毛藻(Chaetoceros compressus)、洛氏角毛藻(Chaetoceros lorenzianus)以及海洋角毛藻(Chaetoceros pelagicus),各优势种的分布趋势相似,呈现出近岸高于外海的趋势。Shannon-Weiner多样性指数与Pielou均匀度指数的平均值分别为2.37和0.56。浮游植物昼夜垂直变化明显。典范对应分析结果显示浮游植物空间分布主要受总氮盐、硅酸盐、微型浮游动物与盐度的影响。根据细胞丰度进行的站位的聚类分析显示浮游植物群落明显可划分为近岸群聚与大洋群聚。