Copepod hatching success in marine ecosystems with high diatom concentrations

Diatoms dominate spring bloom phytoplankton assemblages in temperate waters and coastal upwelling regions of the global ocean. Copepods usually dominate the zooplankton in these regions and are the prey of many larval fish species. Recent laboratory studies suggest that diatoms may have a deleterious effect on the success of copepod egg hatching. These findings challenge the classical view of marine food-web energy flow from diatoms to fish by means of copepods. Egg mortality is an important factor in copepod population dynamics, thus, if diatoms have a deleterious in situ effect, paradoxically, high diatom abundance could limit secondary production. Therefore, the current understanding of energy transfer from primary production to fisheries in some of the most productive and economically important marine ecosystems may be seriously flawed. Here we present in situ estimates of copepod egg hatching success from twelve globally distributed areas, where diatoms dominate the phytoplankton assemblage. We did not observe a negative relationship between copepod egg hatching success and either diatom biomass or dominance in the microplankton in any of these regions. The classical model for diatom-dominated system remains valid.     

Vast laminated diatom mat deposits from the west low-latitude Pacific Ocean in the last glacial period

Diatoms are one of the predominant contributors to global carbon fixation by accounting for over 40% of total oceanic primary production and dominate export production. They play a significant role in marine biogeochemistry cycle. The diatom mat deposits are results of vast diatoms bloom. By analysis of diatom mats in 136°00′--140°00′E, 15°00′--21°00′N, Eastern Philippines Sea, we identified the species of the diatoms as giant Ethmodiscus rex （Wallich） Hendey. AMS 14C dating shows that the sediments rich in diatom mats occurred during 16000--28600 a B.P., which means the bloom mainly occurred during the last glacial period, while there are no diatom mat deposits in other layers. Preliminary analysis indicates that Antarctic Intermediate Water （AAIW） expanded northward and brought silicate-rich water into the area, namely, silicon leakage processes caused the bloom of diatoms. In addition, the increase of iron input is one of the main reasons for the diatom bloom.     

Photosynthetic architecture differs in coastal and oceanic diatoms

Diatoms are a key taxon of eukaryotic phytoplankton and a major contributor to global carbon fixation. They are ubiquitous in the marine ecosystem despite marked gradients in environmental properties, such as dissolved iron concentrations, between coastal and oceanic waters. Previous studies have shown that offshore species of diatoms and other eukaryotic algae have evolved lower iron requirements to subsist in iron-poor oceanic waters, but the biochemical mechanisms responsible for their decreased iron demand are unknown. Here we show, using laboratory-cultured model species, a fundamental difference between a coastal and an oceanic diatom in their photosynthetic architecture. Specifically, the oceanic diatom had up to fivefold lower photosystem I and up to sevenfold lower cytochrome b6f complex concentrations than a coastal diatom. These changes to the photosynthetic apparatus markedly decrease the cellular iron requirements of the oceanic diatom but not its photosynthetic rates. However, oceanic diatoms might have also sacrificed their ability to acclimate to rapid fluctuations in light intensity--a characteristic of dynamic and turbid coastal waters. We suggest that diatoms, and probably other eukaryotic algal taxa, exploited this difference in the underwater light climate between oceanic and coastal waters, enabling them to decrease their iron requirements without compromising photosynthetic capacity. This adaptation probably facilitated the colonization of the open ocean by diatoms, and contributes to their persistence in this iron-impoverished environment.     

Architecture and material properties of diatom shells provide effective mechanical protection

Diatoms are the major contributors to phytoplankton blooms in lakes and in the sea and hence are central in aquatic ecosystems and the global carbon cycle. All free-living diatoms differ from other phytoplankton groups in having silicified cell walls in the form of two 'shells' (the frustule) of manifold shape and intricate architecture whose function and role, if any, in contributing to the evolutionary success of diatoms is under debate. We explored the defence potential of the frustules as armour against predators by measuring their strength. Real and virtual loading tests (using calibrated glass microneedles and finite element analysis) were performed on centric and pennate diatom cells. Here we show that the frustules are remarkably strong by virtue of their architecture and the material properties of the diatom silica. We conclude that diatom frustules have evolved as mechanical protection for the cells because exceptional force is required to break them. The evolutionary arms race between diatoms and their specialized predators will have had considerable influence in structuring pelagic food webs and biogeochemical cycles.     

The decline and fate of an iron-induced subarctic phytoplankton bloom

Iron supply has a key role in stimulating phytoplankton blooms in high-nitrate low-chlorophyll oceanic waters. However, the fate of the carbon fixed by these blooms, and how efficiently it is exported into the ocean's interior, remains largely unknown. Here we report on the decline and fate of an iron-stimulated diatom bloom in the Gulf of Alaska. The bloom terminated on day 18, following the depletion of iron and then silicic acid, after which mixed-layer particulate organic carbon (POC) concentrations declined over six days. Increased particulate silica export via sinking diatoms was recorded in sediment traps at depths between 50 and 125 m from day 21, yet increased POC export was not evident until day 24. Only a small proportion of the mixed-layer POC was intercepted by the traps, with more than half of the mixed-layer POC deficit attributable to bacterial remineralization and mesozooplankton grazing. The depletion of silicic acid and the inefficient transfer of iron-increased POC below the permanent thermocline have major implications both for the biogeochemical interpretation of times of greater iron supply in the geological past, and also for proposed geo-engineering schemes to increase oceanic carbon sequestration.     

Importance of stirring in the development of an iron-fertilized phytoplankton bloom

The growth of populations is known to be influenced by dispersal, which has often been described as purely diffusive. In the open ocean, however, the tendrils and filaments of phytoplankton populations provide evidence for dispersal by stirring. Despite the apparent importance of horizontal stirring for plankton ecology, this process remains poorly characterized. Here we investigate the development of a discrete phytoplankton bloom, which was initiated by the iron fertilization of a patch of water (7 km in diameter) in the Southern Ocean. Satellite images show a striking, 150-km-long bloom near the experimental site, six weeks after the initial fertilization. We argue that the ribbon-like bloom was produced from the fertilized patch through stirring, growth and diffusion, and we derive an estimate of the stirring rate. In this case, stirring acts as an important control on bloom development, mixing phytoplankton and iron out of the patch, but also entraining silicate. This may have prevented the onset of silicate limitation, and so allowed the bloom to continue for as long as there was sufficient iron. Stirring in the ocean is likely to be variable, so blooms that are initially similar may develop very differently.     

Deep carbon export from a Southern Ocean iron-fertilized diatom bloom

Fertilization of the ocean by adding iron compounds has induced diatom-dominated phytoplankton blooms accompanied by considerable carbon dioxide drawdown in the ocean surface layer. However, because the fate of bloom biomass could not be adequately resolved in these experiments, the timescales of carbon sequestration from the atmosphere are uncertain. Here we report the results of a five-week experiment carried out in the closed core of a vertically coherent, mesoscale eddy of the Antarctic Circumpolar Current, during which we tracked sinking particles from the surface to the deep-sea floor. A large diatom bloom peaked in the fourth week after fertilization. This was followed by mass mortality of several diatom species that formed rapidly sinking, mucilaginous aggregates of entangled cells and chains. Taken together, multiple lines of evidence-although each with important uncertainties-lead us to conclude that at least half the bloom biomass sank far below a depth of 1,000 metres and that a substantial portion is likely to have reached the sea floor. Thus, iron-fertilized diatom blooms may sequester carbon for timescales of centuries in ocean bottom water and for longer in the sediments.     

Evolution and metabolic significance of the urea cycle in photosynthetic diatoms

Diatoms dominate the biomass of phytoplankton in nutrient-rich conditions and form the basis of some of the world's most productive marine food webs. The diatom nuclear genome contains genes with bacterial and plastid origins as well as genes of the secondary endosymbiotic host (the exosymbiont), yet little is known about the relative contribution of each gene group to diatom metabolism. Here we show that the exosymbiont-derived ornithine-urea cycle, which is similar to that of metazoans but is absent in green algae and plants, facilitates rapid recovery from prolonged nitrogen limitation. RNA-interference-mediated knockdown of a mitochondrial carbamoyl phosphate synthase impairs the response of nitrogen-limited diatoms to nitrogen addition. Metabolomic analyses indicate that intermediates in the ornithine-urea cycle are particularly depleted and that both the tricarboxylic acid cycle and the glutamine synthetase/glutamate synthase cycles are linked directly with the ornithine-urea cycle. Several other depleted metabolites are generated from ornithine-urea cycle intermediates by the products of genes laterally acquired from bacteria. This metabolic coupling of bacterial- and exosymbiont-derived proteins seems to be fundamental to diatom physiology because the compounds affected include the major diatom osmolyte proline and the precursors for long-chain polyamines required for silica precipitation during cell wall formation. So far, the ornithine-urea cycle is only known for its essential role in the removal of fixed nitrogen in metazoans. In diatoms, this cycle serves as a distribution and repackaging hub for inorganic carbon and nitrogen and contributes significantly to the metabolic response of diatoms to episodic nitrogen availability. The diatom ornithine-urea cycle therefore represents a key pathway for anaplerotic carbon fixation into nitrogenous compounds that are essential for diatom growth and for the contribution of diatoms to marine productivity.     

夏季南极长城湾沿岸海域浮游硅藻群落定量分析

对南极硅藻进行定量研究较有必要,以客观准确地了解它的组成和分布特征。在南极长城湾沿岸海域共设置6个采样点,每个采样点定量采集3个浮游硅藻样本。结果显示:长城湾海域共发现浮游硅藻31种。浮游硅藻群落的优势种为Cocconesi fasciolata(18.5%)、Amphora holsatica(13.6%)。次优势种为Cocconeis costata(7.1%)、Cocconeis imperatrix(6.8%)、Thalassiosira antarctica(6.4%)、Rhoicosphenia genuflexa(6.1%)、Licmophora antarctica(5.1%)。各样点浮游硅藻细胞密度总平均值为1.92×10~5ind·L~(-1),变幅为1.44×10~5~2.56×10~5ind·L~(-1)。浮游硅藻的分布从湾心到湾口呈现出一定趋势:浮游硅藻的物种丰度,湾心处共15种,湾口处共25种;浮游硅藻的细胞密度,湾心处平均为2.51×10~5ind·L-1,湾口处平均为1.48×10~5ind·L~(-1)。通过相关性分析结果可知,浮游硅藻由湾心到湾口,物种丰度和硅藻细胞密度呈负相关。     

海洋浮游植物休眠期的生态学研究

休眠期是甲藻和硅藻抵御外界不利环境条件的一种有效的生理反应机制，也是许多赤潮藻生活史中的一个重要生活阶段．从生态分布、生态作用尤其是在赤潮生消过程中的作用等角度简述了海洋浮游甲藻和硅藻休眠期的生态学的研究情况，并对二者的生态学特征进行比较．甲藻孢囊和硅藻休止细胞在海洋沉积物中的丰度有明显差别，前者较后者具有更明显的季节性萌发能力．温度和光照是影响二者萌发的重要生态因子，温度是甲藻孢囊萌发不可或缺的因素，而光照是硅藻休止细胞萌发的触发因子．二者在种源策略和生态特征上的差异导致了不同的群落演替模式和引发藻华的机制．     

Unicellular C4 photosynthesis in a marine diatom

Nearly 50 years ago, inorganic carbon was shown to be fixed in microalgae as the C3 compound phosphoglyceric acid. The enzyme responsible for C3 carbon fixation, ribulose-1,5-bisphosphate carboxylase (Rubisco), however, requires inorganic carbon in the form of CO2 (ref. 2), and Rubisco enzymes from diatoms have half-saturation constants for CO2 of 30-60 microM (ref. 3). As a result, diatoms growing in seawater that contains about 10 microM CO2 may be CO2 limited. Kinetic and growth studies have shown that diatoms can avoid CO2 limitation, but the biochemistry of the underlying mechanisms remains unknown. Here we present evidence that C4 photosynthesis supports carbon assimilation in the marine diatom Thalassiosira weissflogii, thus providing a biochemical explanation for CO2-insensitive photosynthesis in marine diatoms. If C4 photosynthesis is common among marine diatoms, it may account for a significant portion of carbon fixation and export in the ocean, and would explain the greater enrichment of 13C in diatoms compared with other classes of phytoplankton. Unicellular C4 carbon assimilation may have predated the appearance of multicellular C4 plants.     

海洋硅藻多样性与生态作用研究

硅藻是海洋浮游植物的主要类群,也是海洋生态系统中的主要初级生产者,在海洋生态系统的物质循环和能量流动中起着极其重要的作用,但人们对海洋硅藻多样性和生态作用的认识还很有限.在叙述海洋硅藻多样性、生态作用及硅藻分类学与系统学的国内外研究动态和最新进展的基础上,概述了厦门大学硅藻课题组近年来在海洋硅藻分类学与系统学、多样性与生态分布、硅藻数据库与计算机自动识别、微型硅藻与硅藻隐形种、硅藻遗传地理差异、硅藻对环境和气候变化的响应机制、硅藻细胞硅吸收与硅质壁形成机制等方面的主要研究进展和取得的主要成果.     

The biogeochemical behavior of dissolved aluminum in the southern Yellow Sea: Influence of the spring phytoplankton bloom

Many recent studies have investigated the nutrient-type profiles of dissolved aluminum(Al) in the ocean.Significant scavenging of dissolved Al can occur during phytoplankton blooms,but the mechanism remains unclear.The distribution of dissolved Al in the southern Yellow Sea(SYS) was investigated in winter and spring 2009.Following measurements at grid stations during the spring sampling cruise,two drifting anchor surveys of more than 100 h were conducted to trace the variation of dissolved Al concentration during the spring phytoplankton bloom(SPB).The concentration of dissolved Al in the SYS decreased from 40 nmol/L in February to 30 nmol/L in March and 10-20 nmol/L in April,while the concentration of Chl a increased from < 2 μg/L in March to > 4 μg/L in April.The concentration of dissolved Al in the SYS decreased significantly with the development of the phytoplankton bloom,which indicated biological scavenging of dissolved Al from water column.The proportion of dissolved Al scavenged from water column by different phytoplankton species differed at the two drifting stations,with greater removal efficiency demonstrated by diatoms than dinoflagellates.Phytoplankton samples collected from the Chl a maximum layer were washed with trace metal clean reagent(oxalate-EDTA-citrate,abbreviate as oxalate solution,Tovar-Sanchez et al.,2003) to enable the surface-scavenged(extracellular) and intracellular Al pools associated with phytoplankton to be differentiated.Thirty-nine to ninetysix percent of the total Al was found to be existed in the interior pools,which indicated that biological absorption was the important way to scavenge dissolved Al during phytoplankton blooms in the SYS.     

The Phaeodactylum genome reveals the evolutionary history of diatom genomes

Diatoms are photosynthetic secondary endosymbionts found throughout marine and freshwater environments, and are believed to be responsible for around one-fifth of the primary productivity on Earth. The genome sequence of the marine centric diatom Thalassiosira pseudonana was recently reported, revealing a wealth of information about diatom biology. Here we report the complete genome sequence of the pennate diatom Phaeodactylum tricornutum and compare it with that of T. pseudonana to clarify evolutionary origins, functional significance and ubiquity of these features throughout diatoms. In spite of the fact that the pennate and centric lineages have only been diverging for 90 million years, their genome structures are dramatically different and a substantial fraction of genes ( approximately 40%) are not shared by these representatives of the two lineages. Analysis of molecular divergence compared with yeasts and metazoans reveals rapid rates of gene diversification in diatoms. Contributing factors include selective gene family expansions, differential losses and gains of genes and introns, and differential mobilization of transposable elements. Most significantly, we document the presence of hundreds of genes from bacteria. More than 300 of these gene transfers are found in both diatoms, attesting to their ancient origins, and many are likely to provide novel possibilities for metabolite management and for perception of environmental signals. These findings go a long way towards explaining the incredible diversity and success of the diatoms in contemporary oceans.     

渤海天津海域的网采浮游植物群落结构与赤潮植物的初步研究

2002年5月-2003年9月对渤海天津海域赤潮监控区进行了网采浮游植物的生态学调查，天津海域共发现浮游植物37属68种，其中硅藻为优势类群，共24属51种，占总种数的75．00％；甲藻7属11种；其他6属6种为少量的蓝藻、黄藻和金藻，其生态类型主要为温带近岸型。圆筛藻属的种类是该海域的全年优势类群，在群落总细胞数中的比例平均高达63．41%。浮游植物群落以大型细胞硅藻．甲藻占优群落为主，其平面分布与环境因子密切相关。     

A mesoscale phytoplankton bloom in the polar Southern Ocean stimulated by iron fertilization

Changes in iron supply to oceanic plankton are thought to have a significant effect on concentrations of atmospheric carbon dioxide by altering rates of carbon sequestration, a theory known as the 'iron hypothesis'. For this reason, it is important to understand the response of pelagic biota to increased iron supply. Here we report the results of a mesoscale iron fertilization experiment in the polar Southern Ocean, where the potential to sequester iron-elevated algal carbon is probably greatest. Increased iron supply led to elevated phytoplankton biomass and rates of photosynthesis in surface waters, causing a large drawdown of carbon dioxide and macronutrients, and elevated dimethyl sulphide levels after 13 days. This drawdown was mostly due to the proliferation of diatom stocks. But downward export of biogenic carbon was not increased. Moreover, satellite observations of this massive bloom 30 days later, suggest that a sufficient proportion of the added iron was retained in surface waters. Our findings demonstrate that iron supply controls phytoplankton growth and community composition during summer in these polar Southern Ocean waters, but the fate of algal carbon remains unknown and depends on the interplay between the processes controlling export, remineralisation and timescales of water mass subduction.     

A massive phytoplankton bloom induced by an ecosystem-scale iron fertilization experiment in the equatorial Pacific Ocean

The seeding of an expanse of surface waters in the equatorial Pacific Ocean with low concentrations of dissolved iron triggered a massive phytoplankton bloom which consumed large quantities of carbon dioxide and nitrate that these microscopic plants cannot fully utilize under natural conditions. These and other observations provide unequivocal support for the hypothesis that phytoplankton growth in this oceanic region is limited by iron bioavailability.     

Mesoscale vertical motion and the size structure of phytoplankton in the ocean

Phytoplankton size structure is acknowledged as a fundamental property determining energy flow through 'microbial' or 'herbivore' pathways. The balance between these two pathways determines the ability of the ecosystem to recycle carbon within the upper layer or to export it to the ocean interior. Small cells are usually characteristic of oligotrophic, stratified ocean waters, in which regenerated ammonium is the only available form of inorganic nitrogen and recycling dominates. Large cells seem to characterize phytoplankton in which inputs of nitrate enter the euphotic layer and exported production is higher. But the size structure of phytoplankton may depend more directly on hydrodynamical forces than on the source of available nitrogen. Here we present an empirical model that relates the magnitude of mesoscale vertical motion to the slope of the size-abundance spectrum of phytoplankton in a frontal ecosystem. Our model indicates that the relative proportion of large cells increases with the magnitude of the upward velocity. This suggests that mesoscale vertical motion-a ubiquitous feature of eddies and unstable fronts-controls directly the size structure of phytoplankton in the ocean.     

广西金鸡滩库区水深梯度下的硅藻生态特征研究

基于生物群落结构的生态水文特征,特别是水深梯度下的硅藻生物群落差异为研究历史水位变迁提供了重要依据,亦为水质生物监测的合理采样水深提供参考意见.通过采集金鸡滩库区不同水深下的附生硅藻,研究了喀斯特地区水深梯度下的硅藻分布特征,探讨了金鸡滩库区硅藻种属对水深变化的响应.研究结果表明曲壳藻属是该库区的绝对优势种,在2.2m深处达到峰值;在水深梯度上,桥弯藻属和脆杆藻属的丰度减小,异极藻属和直链藻属的丰度增加.硅藻的生态类群特征为:硅藻种群的耐污性和N-异氧类群增加,营养偏好和pH偏好呈波动变化,盐度偏好变化不明显.在不同水深下,各指数的相对离散程度很低,认为在合理水深内的生物采样对水质生物评价影响较小;在响应水深梯度上,SHE是所有生物指数中较理想的.该研究结果为重塑本地区的古水位、古洪水提供了重要的理论依据,对硅藻生物监测水质具有一定的指导意义.     

2010年夏季广西北部湾沿岸浮游植物分布特征及其与环境因子的相关性

2010年6月4日至26日在广西北部湾沿岸共设46个站位凋查分析浮游植物群落结构特征,以及浮游植物数量与环境因子的相关性。结果共鉴定出浮游植物70属149种(含变种和变型),其中硅藻41属106种,占总物种数的71.14%,占总细胞丰度的95.81%;甲藻18属32种,占总物种数的21.48%,占总细胞丰度的1.44%。浮游植物平均密度为61.10×104 cell/L,硅藻平均细胞丰度为58.54×104 cell/L,甲藻类平均细胞丰度为0.88×104cell/L。浮游植物密集区在防城港海区,此外在北海营盘附近海域有一个较高的密度区。浮游植物丰度在茅尾海和钦州港近岸较低,其余海区的浮游植物数量分布态势为近岸高、外海低。浮游植物主要优势种为尖刺拟菱形藻(Pseudo-nitzschia pungens)、中肋骨条藻(Skeletonema costatum)、拟旋链角毛藻(Chaetocerospseudocurvisetus)、菱形海线藻(Thalassionema nitzschioides)、丹麦细柱藻(Leptocylindrus danicus)。尖刺拟菱形藻、拟旋链角毛藻密集区主要在防城港海区,而中肋骨条藻密集区主要在北海海区。浮游植物数量与温度、盐度及硝酸盐、亚硝酸盐、氨盐和硅酸盐无相关性,北海海区和防城港海区浮游植物丰度与活性磷酸盐呈显著正相关,磷在以上两个海区成为浮游植物生长的限制因子。