硅藻硅酸转运子（silicon transporters, SITs）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。硅酸转运子(silicon transporters, SITs)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SITs的分子特征、分子机制、功能多样性及进化关系,为SITs转运机制的研究提供了理论基础。     

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。     

海洋硅藻硅质细胞壁结构的形成机理研究概述

硅藻具有形态各异、结构复杂、精美绝伦的硅质细胞壁,是海洋中进行生物硅化最主要的生物体。硅质细胞壁的形成同样是一个错综复杂的过程,它涉及硅藻细胞将硅酸从胞外转运到胞内;硅酸在细胞内的转移;在硅沉积囊泡(SDV)中的浓缩沉积;最后合成具种类特异性的细胞壁。重点介绍硅酸转运基因(SITs)的分子特征与作用机制;与生物硅化相关的三种蛋白即硅体蛋白(frus-tulins)、亲硅蛋白(silaffins)和侧壁蛋白(pleuralins)的结构与功能;硅质结构如何在硅沉积囊泡内最终形成的模式。     

Fronts and surface zonal geostrophic current along 115°E in the southern Indian Ocean

硅藻具有形态各异、结构复杂、精美绝伦的硅质细胞壁,是海洋中进行生物硅化最主要的生物体。硅质细胞壁的形成同样是一个错综复杂的过程,它涉及硅藻细胞将硅酸从胞外转运到胞内;硅酸在细胞内的转移;在硅沉积囊泡(SDV)中的浓缩沉积;最后合成具种类特异性的细胞壁。重点介绍硅酸转运基因(SITs)的分子特征与作用机制;与生物硅化相关的三种蛋白即硅体蛋白(frustulins)、亲硅蛋白(silaffins)和侧壁蛋白(pleuralins)的结构与功能;硅质结构如何在硅沉积囊泡内最终形成的模式。     

海水中可溶性无机硅

海水中可溶性无机硅(以下简称硅),是海洋浮游植物所必须的营养盐之一,尤其是对硅藻类浮游植物、放射虫和硅质海绵,硅更是构成机体不可缺少的组分。在海洋浮游植物中硅藻占很大部分,硅藻繁殖时摄取硅使海水中硅的含量下降。浮游植物死亡下沉分解又释放出硅,使海水中硅的含量再生。另外放射虫也能排泄出二氧化硅,硅质死亡迅速释放二氧化硅进入海水中。总之硅的含量对于海洋生物产量具有直接和间     

第五节 海水中可溶性无机硅

海水巾可溶性无机硅(以下简称硅)，是海洋浮游植物所必须的营养盐之一，尤其是对硅藻类浮游植物、放射虫和硅质海绵，硅更是构成机体不可缺少的组分。在海洋浮游植物中硅藻占很大部分，硅藻繁殖时摄取硅使海水中硅的含量下降。浮游植物死亡下沉分解又释放出硅，使海水中硅的含量再生。另外放射虫也能排泄出二氧化硅，硅质死亡迅速释放二氧化硅进入海水中。总之硅的含量对于海洋生物产量具有直接和间接影响，由此可知生物活动使海水中硅的含量随季节呈现周期性变化。     

悬浮颗粒物中生物硅测定方法的改进与应用

硅是河流和海洋中非常重要的营养盐,硅藻、放射虫、硅质海绵、硅鞭毛虫的生长和骨骼形成都离不开硅[1].硅在河流和沿岸海域生态系统中扮演着非常重要的角色,在浮游植物大量繁殖季节,由于硅藻的大量繁殖,使活性硅酸盐含量急剧下降,甚至使硅藻的生长繁殖受到限制.在其他环境条件适宜的情况下,充足的硅又将成为发生硅藻赤潮的物质基础[2].生物硅是用化学方法测定的无定形硅的含量[3],称为生物蛋白石或简称蛋白石,是地球化学、古海洋学研究中非常重要的参数[4].根据世界部分河流生物硅和溶解硅的平均值计算,河流输送入海的硅中有16%的是生物硅,因此河流输送的生物硅的量是全球海洋硅收支中不可忽略的部分[5].     

南海南部表层沉积物中生物硅的分布及其环境意义

对南海南部25个表层沉积样进行了生物硅的测定分析,试图揭示南海南部表层沉积生物硅的分布及其对现代海洋环境的指示意义,以便为古海洋学研究提供进一步的科学依据。研究发现,表层沉积物中生物硅含量与其所处水深呈显著正相关关系,相关系数达到0．782。陆架浅水区表层沉积物中生物硅含量非常低,不能反映表层水体中硅质生物生产力情况,这可能与沉积类型和陆源物质输入影响有关。深水区表层沉积物中生物硅的含量分布表明,其不仅能反映出表层水体中硅质生物的古生产力水平,而且还能指示上升流的强弱,从而进一步证实了利用沉积物中生物硅含量来追踪上升流发育和变化的有效性与可信度。研究结果还显示,在研究区域中北部表层沉积生物硅中放射虫和海绵骨针较硅藻占有更大的比重,这可能是由于硅藻易被溶解并易被其他生物体摄食的缘故。在有上升流发育的海域,放射虫、硅藻和海绵骨针基本上均表现出较高的丰度,这与高的生物硅含量相一致。     

南海南部表层沉积物中生物硅的分布及其环境意义

对南海南部25个表层沉积样进行了生物硅的测定分析,试图揭示南海南部表层沉积生物硅的分布及其对现代海洋环境的指示意义,以便为古海洋学研究提供进一步的科学依据。研究发现,表层沉积物中生物硅含量与其所处水深呈显著正相关关系,相关系数达到0.782。陆架浅水区表层沉积物中生物硅含量非常低,不能反映表层水体中硅质生物生产力情况,这可能与沉积类型和陆源物质输入影响有关。深水区表层沉积物中生物硅的含量分布表明,其不仅能反映表层水体中硅质生物的古生产力水平,而且还能指示上升流的强弱,从而进一步证实了利用沉积物中生物硅含量来追踪上升流发育和变化的有效性与可信度。研究结果还显示,在研究区域中北部表层沉积生物硅中放射虫和海绵骨针较硅藻占有更大的比重,这可能是由于硅藻易被溶解并易被其他生物体摄食的缘故。在有上升流发育的海域,放射虫、硅藻和海绵骨针基本上表现出较高的丰度,这与高的生物硅含量相一致。     

沉积物中生物硅分析方法评述

硅是水生态系统中构成生物群落的重要元素。硅藻、放射虫、硅质海绵和硅鞭毛虫生长和骨骼形成都离不开硅。ＤｅＭａｓｔｅｒ１９８１年研究指出海洋硅主要来源于河流(４．２±０．８×１０１４ｇ／ａ，ＳｉＯ２)和热液喷发 (１．９±１．０×１０１４ｇ／ａ，ＳｉＯ２)。另外冰川风化、低温海底玄武岩风化和沉积物的成岩作用亦是其来源之一。而Ｃｏｎｌｅｙ等１９９３年报道富营养化往往引起水体中溶解硅的浓度逐渐降低。生物硅（Ｂｉｏｇｅｎｉｃｓｉｌｉｃａ）指化学方法测定的无定形硅的含量 ,亦称为生物蛋白石或简称蛋白石。生源无定形…     

Bioavailability of different chemical forms of dissolved silica can affect marine diatom growth

In this study, we demonstrate that dissolved silica obtained from mineral (crystalline quartz), biogenic amorphous (diatomaceous earth) and artificial amorphous sources (Aerosil) influence the growth rate of two marine diatoms, Chaetoceros sp. and Skeletonema marinoi. Diatoms were reared in four different experimental conditions in artificial seawater containing either dissolved silica previously obtained through dissolution of the mineral crystalline quartz or two amorphous substrates, biogenic diatomaceous earth or artificial Aerosil silica. Sodium metasilicate was used as control. When the silica in the different media reached concentrations higher than 107 μm , particles were eliminated by filtration and the diatom cells were inoculated. Maximum cell density, growth and silica assimilation rates of both species in the presence of dissolved silica derived from crystalline quartz and metasilicate were higher than those obtained with the other silica sources. These results are discussed against the background of previous geochemical studies that have shown that silica–water interactions are strictly dependent on the silica polymorphs involved and on the ionic composition of the solution. Our results demonstrate that the soluble silicon compounds generated in seawater by crystalline sources are highly bioavailable compared with those generated by biogenic and amorphous materials. These findings are potentially of considerable ecological importance and may contribute to clarifying anomalous spatial and temporal distributions of siliceous organisms with respect to the presence of lithogenic or biogenic silica sources in marine environments.     

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.     

Particle sedimentation patterns in the eastern Fram Strait during 2000–2005: Results from the Arctic long-term observatory HAUSGARTEN

Since 2000 long-term measurements of vertical particle flux have been performed with moored sediment traps at the long-term observatory HAUSGARTEN in the eastern Fram Strait (79°N/4°E). The study area, which is seasonally covered with ice, is located in the confluence zone of the northward flowing warm saline Atlantic water with cold, low salinity water masses of Arctic origin. Current projections suggest that this area is particularly vulnerable to global warming. Total matter fluxes and components thereof (carbonate, particulate organic carbon and nitrogen, biogenic silica, biomarkers) revealed a bimodal seasonal pattern showing elevated sedimentation rates during May/June and August/September. Annual total matter flux (dry weight, DW) at ～300 m depth varied between 13 and 32 g m^?2 a^?1 during 2000 and 2005. Of this total flux 6–13% was due to CaCO₃, 4–21% to refractory particulate organic carbon (POC), and 3–8% to biogenic particulate silica (bPSi). The annual flux of all biogenic components together was almost constant during the period studied (8.5–8.8 g m^?2 a^?1), although this varied from 27% to 67% of the total annual flux. The fraction was lowest in a year characterized by the longest duration of ice coverage (91 and 70 days for the calendar year and summer season, May–September, respectively). Biomarker analyses revealed that organic matter originating from marine sources was present in excess of terrigenious material in the sedimented matter throughout most of the study period. Fluxes of recognizable phyto- and protozooplankton cells amounted up to 60×10⁶ m^?2 d^?1. Diatoms and coccolithophorids were the most abundant organisms. Diatoms, mainly pennate species, dominated during the first years of the investigation. A shift in the composition occurred during the last year when numbers of diatoms declined considerably, leading to a dominance of coccolithoporids. This was also reflected in a decrease in the sedimentation of bPSi. The sedimentation of biogenic matter, however, did not differ from the amount observed during the previous years. Among the larger organisms, pteropods at times contributed significantly to both the total matter and CaCO₃, fluxes.     

Modeling the long-term variability of phytoplankton functional groups and primary productivity in the South China Sea

Primary productivity (PP) and phytoplankton structure play an important role in regulating oceanic carbon cycle. The unique seasonal circulation and upwelling pattern of the South China Sea (SCS) provide an ideal natural laboratory to study the response of nutrients and phytoplankton dynamics to climate variation. In this study, we used a three-dimensional (3D) physical–biogeochemical coupled model to simulate nutrients, phytoplankton biomass, PP, and functional groups in the SCS from 1958 to 2009. The modeled results showed that the annual mean carbon composition of small phytoplankton, diatoms, and coccolithophores was 33.7, 52.7, and 13.6 %, respectively. Diatoms showed a higher seasonal variability than small phytoplankton and coccolithophores. Diatoms were abundant during winter in most areas of the SCS except for the offshore of southeastern Vietnam, where diatom blooms occurred in both summer and winter. Higher values of small phytoplankton and coccolithophores occurred mostly in summer. Our modeled results indicated that the seasonal variability of PP was driven by the East Asian Monsoon. The northeast winter monsoon results in more nutrients in the offshore area of the northwestern Luzon Island and the Sunda Shelf, while the southwest summer monsoon drives coastal upwelling to bring sufficient nutrients to the offshore area of southeastern Vietnam. The modeled PP was correlated with El Niño/Southern Oscillation (ENSO) at the interannual scale. The positive phase of ENSO (El Niño conditions) corresponded to lower PP and the negative phase of ENSO (La Niña conditions) corresponded to higher PP.     

Winter distribution of diatom assemblages along the coastline of R. O. Korea in 2010

Diatoms are a globally successful and eukaryotic photosynthetic organism with an ornamented silica external wall. The relationship between their valve morphology and habitat means that diatoms can be used as bioindicators to characterize the aquatic environment. To estimate the differential distribution and diversity of diatom assemblages along the coastal line, we collected phytoplankton samples from 114 coastal sites of waters of R. O. Korea. We applied the unweighted pair-group technique usin...     

The annual silica cycle of the North Pacific subtropical gyre

Silica cycling in the upper 175 m of the North Pacific Subtropical Gyre was examined over a two year period (January 2008-December 2009) at the Hawaii Ocean Time-series (HOT) station ALOHA. Silicic acid concentrations in surface waters ranged from 0.6 to 1.6 ??M, exhibiting no clear seasonal trends. Biogenic silica concentrations and silica production rates increased by an order of magnitude each summer following stratification of the upper 50 m reaching values of 157 nmol Si L⁻¹ and 81 nmol Si L⁻¹ d⁻¹, in 2008 and 2009, respectively. Sea surface height anomalies together with analyses of variability in isothermal surfaces at 150-175 m indicated that the summer periods of elevated biogenic silica were associated with anticyclonic mesoscale features during both years. Lithogenic silica concentrations increased in the spring during the known period of maximum atmospheric dust concentrations with maximum values of 36 nmol Si L⁻¹ in the upper 10 m. Dust deposition would enhance levels of dissolved iron in surface waters, but there was no response of diatom biomass or silica production to increases in near-surface ocean lithogenic silica concentrations suggesting iron sufficiency of diatom silica production rates.Low ambient silicic acid concentrations restricted silica production rates to an average of 43% of maximum potential rates. Si sufficiency only occurred during the summer period when diatom biomass was elevated suggesting that bloom diatoms are adapted to exploit low silicic acid concentrations. Annual silica production at HOT is estimated to be 63 mmol Si m⁻² a⁻¹ with summer blooms contributing 29% of the annual total. Diatoms are estimated to account for 3-7% of total phytoplankton primary productivity, but 9-20% of organic carbon export confirming past suggestions that diatoms are relatively minor contributors to primary productivity and autotrophic biomass, but important contributors to new and export production in oligotrophic open-ocean ecosystems.Annual silica production at HOT is nearly 4-fold lower than estimates at the Bermuda Atlantic Time-series Study (BATS) site in the Sargasso Sea from the 1990s, but annual silica export at the base of the euphotic zone is similar between the two gyres indicating very different balances between silica production and its loss in surface waters. On a relative basis, BATS is a more productive system with respect to silica, where biogenic silica is recycled with high efficiency in surface waters; in contrast the NPSG is a lower productivity system with respect to silica, but where lower recycling efficiency leads to a much higher fraction of new silica production. The two gyres show contrasting long-term trends in diatom biomass as biogenic silica concentrations at HOT have been increasing since 1997, but they have been decreasing at BATS suggesting very different forcing of decadal trends in the contribution of diatoms in carbon cycling between these gyres. Combining the data from both gyres indicates that globally subtropical gyres produce 13 Tmol Si a⁻¹, which is only 51% of previous estimates reducing the contribution of subtropical gyres to 5-7% of global annual marine silica production.     

Deep ocean fluxes and their link to surface ocean processes and the biological pump

Intense studies of upper and deep ocean processes were carried out in the Northwestern Indian Ocean (Arabian Sea) within the framework of JGOFS and related projects in order to improve our understanding of the marine carbon cycle and the ocean’s role as a reservoir for atmospheric CO₂. The results show a pronounced monsoon-driven seasonality with enhanced organic carbon fluxes into the deep-sea during the SW Monsoon and during the early and late NE Monsoon north of 10°N. The productivity is mainly regulated by inputs of nutrients from subsurface waters into the euphotic zone via upwelling and mixed layer-deepening. Deep mixing introduces light limitation by carrying photoautotrophic organisms below the euphotic zone during the peak of the NE Monsoon. Nevertheless, deep mixing and strong upwelling during the SW Monsoon provide an ecological advantage for diatoms over other photoautotrophic organisms by increasing the silica concentrations in the euphotic zone. When silica concentrations fall below 2 μmol l⁻¹, diatoms lose their dominance in the plankton community. During diatom-dominated blooms, the biological pathway of uptake of CO₂ (the biological pump) appears to be more efficient than during blooms of other organisms, as indicated by organic carbon to carbonate carbon (rain) ratios. Due to the seasonal alternation of diatom and non-diatom dominated exports, spatial variations of the annual mean rain ratios are hardly discernible along the main JGOFS transect.Data-based estimates of the annual mean impact of the biological pump on the fCO₂ in the surface water suggest that the biological pump reduces the increase of fCO₂ in the surface water caused by intrusion of CO₂-enriched subsurface water by 50–70%. The remaining 30 to 50% are attributed to CO₂ emissions into the atmosphere. Rain ratios up to 60% higher in river-influenced areas off Pakistan and in the Bay of Bengal than in the open Arabian Sea imply that riverine silica inputs can further enhance the impact of the biological pump on the fCO₂ in the surface water by supporting diatom blooms. Consequently, it is assumed that reduced river discharges caused by the damming of major rivers increase CO₂ emission by lowering silica inputs to the Arabian Sea; this mechanism probably operates in other regions of the world ocean also.     

Effects of virus infection on expression of cell cycle regulatory proteins in the unicellular marine algae Emiliania huxleyi

The authors have investigated the biochemical events by which marine algal virus infection induces cell cycle arrest. The key G₂/M-phase regulatory proteins are analyzed by immunobloting in unicellular Emiliania huxleyi, suggesting that virus induced cell cycle arrest is related with virus''s effect on cyclins and cyclin dependent kinases. E. huxleyi virus (EhV) represses Cdc2/cyclinB complex activity by inhibiting the activity of Cdc2 kinase in a phosphorylation-related manner, blocking host cells G₂/M checkpoint. Dephosphorylated/inactive Cdc25C combined with up-regulation of Wee1 expression at early infect period appears to be important mechanisms by which EhV represses Cdc2/cyclinB complex activity that is required for entry into M phase. This study has allowed us to confirm that algal virus infection leads to selective activation or inhibition of certain cell-cycle factors, which may play a significant role in establishing a more efficient environment for viral gene expression and DNA replication.