海洋中的凝集网与透明胞外聚合颗粒物

透明胞外聚合颗粒物(TEP)是海洋中大量存在的能被爱尔新蓝(alcian blue)染色的由酸性多糖组成的透明胶状颗粒物,主要来源于浮游植物,由于其透明的特性而被长期忽视。TEP同时具有胶体和颗粒物的特性。作为胶体,TEP为细菌提供了栖息场所与降解基质,同时TEP可以吸收痕量元素,以改变这些元素的生物地球化学过程。作为颗粒物,TEP可以聚集并沉降,由于其高的碳含量,会在很大程度上影响海洋的碳通量。由于TEP可以被中型浮游动物所摄食,所以TEP可以连接并缩短微食物环和经典食物链,在海洋生态系统中起很重要的作用。介绍了TEP的定义、测量方法、来源、形成、及其与浮游植物的关系和其生态功能。     

铁施肥促进海洋浮游植物和生物碳泵的不确定性

铁作为浮游植物所必需的微量元素,限制了全球超过三分之一海域的初级生产力,尤其是在高营养盐、低叶绿素海域(high nutrient low chlorophyll,HNLC)。长期以来海洋铁施肥被认为是一项可以降低大气二氧化碳含量的地球工程策略。然而通过13次海洋人工铁施肥(artificial ocean iron fertilization,aOIF)实验发现,铁的额外添加对海洋深层碳输出量的促进作用要显著低于预期。本文简要地总结了碳在海洋和大气中的循环过程,回顾了人工铁施肥实验对生物碳泵和碳通量等的影响,分析了从海洋铁施肥到海洋碳汇关键生物地球化学过程的影响因素。综上分析发现,科学界对生物碳泵过程及其调控机制的认识仍十分浅薄,考虑到海洋铁施肥还会对海洋生态系统带来一定的负面作用,铁施肥能否作为降低大气中CO₂的有效手段,以达到碳中和并缓解温室效应仍需进一步研究。     

全球CO2 水平升高对浮游植物生理和生态影响的研究进展

工业革命以来由于化石燃料的大量燃烧,大气CO₂水平不断增加,预计在21世纪末将增至现有水平的两倍,达到750 μL/L。作为全球初级生产力的重要贡献者,浮游植物应对CO₂水平升高的生理生态响应必然会对水生生态系统和碳、氮等元素的生物地球化学循环产生重要影响。全球CO₂水平的升高将显著改变水体的碳化学环境,淡水生态系统(湖泊和河流)由于容量小变化比海洋更为显著。水体碳化学环境的改变首先会影响浮游植物个体,在高CO₂水平下,浮游植物的细胞会有变小的趋势,并且细胞的光合作用强度也会有不同程度的增加,其中细胞较小或者不具有碳浓缩机制(CCM)的浮游植物增加较多,此外浮游植物细胞的化学元素计量值也将显著改变。随后浮游植物个体水平上的变化会进一步影响水生生态系统,例如水体初级生产力水平的提高,浮游植物、浮游动物群落结构组成以及水体微食物网结构的变化等。此外浮游植物对CO₂水平升高的生理生态响应程度还与水体的营养水平有关。总结了大气CO₂水平升高对浮游植物生理生态影响的研究方法,展望了未来可能的研究方向。     

单细胞原生生物在海洋碳汇研究中的重要性和展望

海洋是地球上最大的碳库,通过对CO~2的固定以及与大气物质和能量的交换,海洋对全球气候的变化起到关键的调控作用。随着全球气候变化的加剧,增加海洋碳汇已经成为应对全球气候变化的热门研究课题和主要途径之一。海洋微型生物在海洋的固碳过程及碳循环中起到关键的作用,对海洋碳汇意义重大。本文综述了一类重要的海洋微型生物——单细胞原生生物在海洋碳汇研究中的重要性,分析了其中的代表——网粘菌门(Labyrintholomycota)原生生物在海洋碳循环和次级生产中的意义,并从清楚地认识海洋碳汇的过程和机制方面,提出未来该领域急需解决的科学问题和可能的研究方案,为丰富海洋碳汇研究的生物学基础提供理论依据。     

采用遥感手段估算海洋初级生产力研究进展

海洋初级生产力的精确估算对渔业资源评估与管理、海洋生态系统和全球变化等研究具有重要意义.传统的现场测量与估算方法必须依赖于随船采样数据.卫星遥感具有能够获取实时的、大尺度的、动态的海洋环境参数的优点,因此卫星遥感日益成为大尺度海洋初级生产力估算的重要手段.本文从海洋水色传感器的发展历程出发,着重归纳了以叶绿素、浮游植物碳和浮游植物吸收系数为参量的海洋初级生产力的遥感估算方法,并就这3类模型的适应性和复杂程度进行了讨论.在此基础上,进一步分析评价了全球海洋初级生产力遥感估算的研究现状.鉴于当前海洋初级生产力遥感估算研究中存在的问题,今后的研究需要在4个方面进一步加强:1)对全球海洋初级生产力估算进行分区域研究;2)加深对浮游植物吸收系数的研究;3)提高海洋遥感技术水平;4)加强实地测量技术的研究.     

典型红树林生态系统藻类多样性及其在生态过程中的作用

藻类是红树林生态系统重要的生物类群, 根据生态习性可分为浮游植物、底栖微藻和大型藻类三个生态类群, 它们在红树林生态系统生物多样性、初级生产、元素循环等方面起着重要作用。但在红树林生态系统中, 关注重点多集中在红树植物和动物, 对其中的藻类重视不够, 且多数研究集中在近20年以及亚洲的红树林区。事实上, 红树林生态系统藻类非常丰富, 其多样性研究有助于深入揭示红树林生态系统的结构与功能。本文介绍了红树林生态系统藻类的组成类群及其重要性, 重点对红树林区浮游植物、底栖硅藻和大型海藻的种类组成、地理分布及其与初级生产力、水质污染、元素循环、碳库形成等生态过程中的作用的研究动态和进展等进行了总结。根据已有研究, 红树林区浮游植物和底栖硅藻的种类数一般为几十到上百种, 其中硅藻在种类和数量上都占绝对优势, 它们是重要的初级生产者、饵料生物和水质污染指示生物; 红树林区底栖大型藻类主要由红藻、绿藻、褐藻、蓝藻组成, 绿藻的种类较多, 红藻在数量上占优势; 藻类是红树林湿地碳库的重要贡献者, 在红树林湿地生态系统碳汇和碳循环中起重要作用。红树林生态系统是个高度动态和异质的系统, 今后应加强红树林藻类多样性的长周期、大尺度变化及不同生境藻类的综合研究, 关注大陆径流和潮汐对藻类多样性和蓝碳的影响, 借助沉积物藻类记录, 探明红树林区藻类的长周期变化, 反演气候变化和人类活动对红树林生态系统的影响过程和机制。     

全球气候变化下的海洋浮游植物多样性

理解全球气候变化对地球生态系统的影响是全世界广泛关注的问题, 而相比于陆地生态系统, 海洋生态系统对全球气候变化更为敏感。全球气候变化对海洋的影响主要表现在海洋暖化、海洋酸化、大洋环流系统的改变、海平面上升、紫外线辐射增强等方面。浮游植物是海洋生态系统最重要的初级生产者, 同时对海洋碳循环起到举足轻重的作用, 其对全球气候变化的响应主要体现在物种分布、初级生产力、群落演替、生物气候学等方面。具体表现在以下方面: 暖水种的分布范围在扩大, 冷水种分布范围在缩小; 浮游植物全球初级生产力降低; 浮游植物群落会向细胞体积更小的物种占优势的方向转变; 浮游植物水华发生的时间提前、强度增强; 一些有害物种水华的发生频率也会增加; 海洋表层海水的酸化会影响浮游植物特别是钙化类群的生长和群落多样性; 紫外辐射增强对浮游植物的生长起到抑制作用; 厄尔尼诺、拉尼娜、降水量的增加通常抑制浮游植物生长。浮游植物生长和分布的变化会体现在多样性的各个层面上。对于浮游植物在全球变化各种驱动因子下的生理生态学和长周期变动观测等是今后研究的重要方向, 也将为理解全球变化下的浮游植物-多样性-生态系统响应与反馈机制提供基本信息。     

基于文献计量的全球海洋酸化研究状况分析

海洋酸化(Ocean acidification)为目前备受人们关注的全球性问题。因此为了能够客观地揭示海洋酸化的研究态势,研究采用文献计量分析(Bibliometric analysis)的方法,以海洋酸化概念提出后(2004年以后)ISI Web of Science期刊引文数据库中涉及到海洋酸化研究的所有文献为样本,对文献的增长趋势及期刊分布进行描述统计,并基于关键词的知识图谱及突变分析的方法探究海洋酸化的热点关注方向随时间的变动及研究前沿。描述统计表明:海洋酸化概念提出的这十多年来,涉及海洋酸化的研究文献数量呈现激增的态势,研究学科交叉明显,海洋酸化对珊瑚礁的影响是这十年来的重点研究领域。从基于关键词的知识图谱可以看到,在海洋酸化研究初期(2004—2009年),研究内容主要分为两个部分,一是海洋酸化对海洋生物(尤其是珊瑚礁生物及浮游植物)及生态系统的影响;二是对海洋酸化现象的认识;中期(2010—2015年),研究内容与初期相似,研究重点往海洋生物上倾斜,同时有新的热点研究区域和研究方向的出现;近期(2016年以后),海洋酸化对海洋生物影响的研究依旧占据着主流研究方向。对基于突变分析得到的当前(2018年2月)海洋酸化研究的热点关注的文献分析发现,当前海洋酸化的研究存在以下5个前沿方向:(1)在探究海洋酸化与生物的关系之时需结合多因子讨论;(2)探索生物在海洋酸化下的内在应对机制;(3)海洋酸化影响下的生物响应的综合评估及预测;(4)探索海洋酸化对海洋生态系统的影响;(5)对海洋酸化概念的挑战——海洋酸化形成原因的探索。     

海洋微型浮游动物对浮游植物和初级生产力的摄食压力

综述了国际上研究微型浮游动物对浮游植物和初级生产力摄食的方法,并重点介绍了稀释法的理论和在实践中遇到的问题。各种方法的微型浮游动物对浮游植物和初级生产力摄食压力的估计表明,微型浮游动物在海洋生态系统中的扮演重要角色。     

基于碳、氮稳定同位素技术的东太湖水生食物网结构

稳定同位素技术是研究生态系统食物网中物质循环与能量流动的有效技术之一。碳稳定同位素比值(δ13C)常用来分析消费者食物来源,而氮稳定同位素比值(δ15N)常用来确定生物在食物网中的营养位置。本研究应用碳、氮稳定同位素技术构建了东太湖食物网结构。结果表明:东太湖食物网主要由两条营养传递途径组成,即浮游植物为初级生产者的浮游营养传递途径和苦草等大型水生植物为初级生产者的近岸底层营养传递途径,湖中9种主要鱼虾类能量主要来自近岸底层传递;翘嘴鲌(Erythroculter ilishaeformis)、鳜(Siniperca chuatsi)和鲶(Silurus sp.)作为湖泊中的顶极捕食者,具有相对最高的营养级,并占据食物网的顶层。     

Diatom-associated bacteria are required for aggregation of Thalassiosira weissflogii

Aggregation of algae, mainly diatoms, is an important process in marine systems leading to the settling of particulate organic carbon predominantly in the form of marine snow. Exudation products of phytoplankton form transparent exopolymer particles (TEP), which acts as the glue for particle aggregation. Heterotrophic bacteria interacting with phytoplankton may influence TEP formation and phytoplankton aggregation. This bacterial impact has not been explored in detail. We hypothesized that bacteria attaching to Thalassiosira weissflogii might interact in a yet-to-be determined manner, which could impact TEP formation and aggregate abundance. The role of individual T. weissflogii-attaching and free-living new bacterial isolates for TEP production and diatom aggregation was investigated in vitro. T. weissflogii did not aggregate in axenic culture, and striking differences in aggregation dynamics and TEP abundance were observed when diatom cultures were inoculated with either diatom-attaching or free-living bacteria. The data indicated that free-living bacteria might not influence aggregation whereas bacteria attaching to diatom cells may increase aggregate formation. Interestingly, photosynthetically inactivated T. weissflogii cells did not aggregate regardless of the presence of bacteria. Comparison of aggregate formation, TEP production, aggregate sinking velocity and solid hydrated density revealed remarkable differences. Both, photosynthetically active T. weissflogii and specific diatom-attaching bacteria were required for aggregation. It was concluded that interactions between heterotrophic bacteria and diatoms increased aggregate formation and particle sinking and thus may enhance the efficiency of the biological pump.     

我国主要海域海水养殖碳汇能力评估及其影响效应——基于我国9个沿海省份面板数据

随着海水养殖业的碳汇功能逐渐被认识和肯定,海水养殖不再单是一项经济活动,而是对环境具有正向影响的碳汇生态活动。以我国沿海9个省份为例,选取海水养殖业碳汇主要贡献的贝类和藻类海产品,并按照各自的碳汇方式对我国沿海地区2008—2015年海水养殖碳汇能力测算,进一步将9个沿海省份按照主要海域划分为渤海、黄海、东海、南海,利用LMDI模型从海水养殖的结构效应和规模效应角度分析碳汇能力的区域差异和主要影响因素。研究结果显示,黄海沿岸海水养殖碳汇能力最强,南海沿岸海水养殖的碳汇转化比例最高,规模效应与我国沿海地区海水养殖碳汇能力始终呈正相关,结构效应的作用显著但不稳定。基于上述结论,我国沿海地区碳汇养殖业应首先提升碳汇养殖技术、稳定海水养殖产量,其次注重优化养殖结构,对碳汇潜力巨大的贝类多加关注。     

中国近海生物固碳强度与潜力

通过现场调查和数据分析,首次探讨了中国近海浮游植物的固碳强度与潜力以及近海人工养殖大型藻类的固碳能力.结果表明,渤、黄、东海的浮游植物固碳强度约为2.22亿t a-1,固碳量的季节变化从大到小的顺序依次为春季、夏季、秋季、冬季,其中春夏季的固碳量占全年的65.3%.南黄海1999～2005年10～11月间浮游植物固碳强度有较大的年际变化,10～11月份7a间其浮游植物最低固碳量为3.54万t d-1,最高为16.58万t d-1,平均为10.50万t d-1,没有明显的年际变化趋势,磷对浮游植物固碳强度的影响最为明显,次之的影响因素是Chl a和亚硝氮 (NO2-N)的含量.南海的固碳能力约为渤黄东海的2倍,为4.16亿t a-1,其季节变化和渤黄东海不同,南海浮游植物在冬季的固碳能力最强,在夏季最弱.整个中国近海浮游植物年固碳量达6.38亿t,可占全球近海区域浮游植物年固碳量的5.77%.实际外海龙须菜的养殖发现,龙须菜每年固定的碳为8.18 t,养殖密度与方式对其产量和固碳量影响巨大.近几年,我国大型经济藻类养殖产量每年在120～150万t左右,换算为固碳量为36～45万t,平均每年40万t,如果海藻养殖产量每年增加5%,到2010年,我国大型经济藻类养殖的固碳量可达57万t a-1,海藻养殖是海洋增加碳汇有多重价值的重要措施.     

Individual and interactive effects of ocean acidification,global warming,and UV radiation on phytoplankton

Rising carbon dioxide (CO₂) concentrations in the atmosphere result in increasing global temperatures and ocean warming (OW). Concomitantly, dissolution of anthropogenic CO₂ declines seawater pH, resulting in ocean acidification (OA) and altering marine chemical environments. The marine biological carbon pump driven by marine photosynthesis plays an important role for oceanic carbon sinks. Therefore, how ocean climate changes affect the amount of carbon fixation by primary producers is closely related to future ocean carbon uptake. OA may upregulate metabolic pathways in phytoplankton, such as upregulating ß-oxidation and the tricarboxylic acid cycle, resulting in increased accumulation of toxic phenolic compounds. Ocean warming decreases global phytoplankton productivity; however, regionally, it may stimulate primary productivity and change phytoplankton community composition, due to different physical and chemical environmental requirements of species. It is still controversial how OA and OW interactively affect marine carbon fixation by photosynthetic organisms. OA impairs the process of calcification in calcifying phytoplankton and aggravate ultraviolet (UV)-induced harms to the cells. Increasing temperatures enhance the activity of cellular repair mechanisms, which mitigates UV-induced damage. The effects of OA, warming, enhanced exposure to UV-B as well as the interactions of these environmental stress factors on phytoplankton productivity and community composition, are discussed in this review.     

Viral lysis of Micromonas pusilla: impacts on dissolved organic matter production and composition

The viral mediated transformation of phytoplankton organic carbon to dissolved forms (“viral shunt”) has been suggested as a major source of dissolved organic carbon (DOC) in marine systems. Despite the potential implications of viral activity on the global carbon fluxes, studies investigating changes in the DOC composition from viral lysis is still lacking. Micromonas pusilla is an ecologically relevant picoeukaryotic phytoplankter, widely distributed in both coastal and oceanic marine waters. Viruses have been found to play a key role in regulating the population dynamics of this species. In this study we used axenic cultures of exponentially growing M. pusilla to determine the impact of viral lysis on the DOC concentration and composition, as estimated from lysate-derived production of transparent exopolymer particles (TEP) and two fractions of fluorescent dissolved organic matter (DOM): aromatic amino acids (excitation/emission; 280/320 nm; F(280/320)) and marine humic-like fluorescent DOM (320/410 nm; F(320/410)). DOC concentration increased 4.5 times faster and reached 2.6 times higher end concentration in the viral infected compared with the non-infected cultures. The production of F(280/320) and F(320/410) were 4.1 and 2.8 times higher in the infected cultures, and the elevated ratio between F(280/320) and F(320/410) in lysates suggested a higher contribution of labile (protein) components in viral produced DOM than in algal exudates. The TEP production was 1.8 times faster and reached a 1.5 times higher level in the viral infected M. pusilla culture compared with the non- infected cultures. The measured increase in both DOC and TEP concentrations suggests that viral lysis has multiple and opposite implications for the production and export processes in the pelagic ocean: (1) by releasing host biomass as DOC it decreases the organic matter sedimentation and promotes respiration and nutrient retention in the photic zone, whereas (2) the observed enhanced TEP production could stimulate particle aggregation and thus carbon export out of the photic zone.     

Mesozooplankton distribution patterns and grazing impacts of copepods and Euphausia crystallorophias in the Amundsen Sea, West Antarctica, during austral summer

The rapid melting of glaciers as well as the loss of sea ice in the Amundsen Sea makes it an ideal environmental setting for the investigation of the impacts of climate change in the Antarctic on the distribution and production of mesozooplankton. We examined the latitudinal distribution of mesozooplankton and their grazing impacts on phytoplankton in the Amundsen Sea during the early austral summer from December 27, 2010 to January 13, 2011. Mesozooplankton followed a latitudinal distribution in relation to hydrographic and environmental features, with copepods dominating in the oceanic area and euphausiids dominating in the polynya. Greater Euphausia crystallorophias biomass in the polynya was associated with lower salinity and higher food concentration (chlorophyll a, choanoflagellates, and heterotrophic dinoflagellates). The grazing impact of three copepods (Rhincalanus gigas, Calanoides acutus, and Metridia gerlachei) on phytoplankton was low, with the consumption of 3 % of phytoplankton standing stock and about 4 % of daily primary production. Estimated daily carbon rations for each of the three copepods were also relatively low (<10 %), barely enough to cover metabolic demands. This suggests that copepods may rely on food other than phytoplankton and that much of the primary production is channeled through microzooplankton. Daily carbon rations for E. crystallorophias were high (up to 49 %) with the grazing impact accounting for 17 % of the phytoplankton biomass and 84 % of primary production. The presence of E. crystallorophias appears to be a critical factor regulating phytoplankton blooms and determining the fate of fixed carbon in the coastal polynyas of the Amundsen Sea.     

Contribution of crenarchaeal autotrophic ammonia oxidizers to the dark primary production in Tyrrhenian deep waters (Central Mediterranean Sea)

Mesophilic Crenarchaeota have recently been thought to be significant contributors to nitrogen (N) and carbon (C) cycling. In this study, we examined the vertical distribution of ammonia-oxidizing Crenarchaeota at offshore site in Southern Tyrrhenian Sea. The median value of the crenachaeal cell to amoA gene ratio was close to one suggesting that virtually all deep-sea Crenarchaeota possess the capacity to oxidize ammonia. Crenarchaea-specific genes, nirK and ureC, for nitrite reductase and urease were identified and their affiliation demonstrated the presence of ‘deep-sea'' clades distinct from ‘shallow'' representatives. Measured deep-sea dark CO₂ fixation estimates were comparable to the median value of photosynthetic biomass production calculated for this area of Tyrrhenian Sea, pointing to the significance of this process in the C cycle of aphotic marine ecosystems. To elucidate the pivotal organisms in this process, we targeted known marine crenarchaeal autotrophy-related genes, coding for acetyl-CoA carboxylase (accA) and 4-hydroxybutyryl-CoA dehydratase (4-hbd). As in case of nirK and ureC, these genes are grouped with deep-sea sequences being distantly related to those retrieved from the epipelagic zone. To pair the molecular data with specific functional attributes we performed [¹⁴C]HCO₃ incorporation experiments followed by analyses of radiolabeled proteins using shotgun proteomics approach. More than 100 oligopeptides were attributed to 40 marine crenarchaeal-specific proteins that are involved in 10 different metabolic processes, including autotrophy. Obtained results provided a clear proof of chemolithoautotrophic physiology of bathypelagic crenarchaeota and indicated that this numerically predominant group of microorganisms facilitate a hitherto unrecognized sink for inorganic C of a global importance.     

海洋生态系统固碳能力估算方法研究进展

气候变化受到全球关注,大气中CO2含量与气候变化息息相关。海洋是地球上最大的活跃碳库,在气候变化中扮演着举足轻重的作用。定量估算海洋中碳元素的吸收、转移、埋藏速率在全球碳循环及全球气候变化研究中有重要意义。目前,海洋固碳能力估算研究包括:利用海-气界面CO2分压差法估算海洋海-气界面CO2交换通量,根据海水中叶绿素含量建立的生态学数理模型法估算真光层浮游生物的初级生产力,234Th—238U不平衡法估算POC输出通量,210Pb定年法估算有机碳沉积通量。但迄今为止的研究工作尚有一定局限性,碳在大气—海水—沉积物3种介质间交换通量间相互影响的研究较少,海洋中碳垂直传输过程的主要影响因素和关键控制因子尚不明确,在海洋生态系统固碳能力估算方法方面国内外还没有统一的规范和标准。为进一步完善海洋生态系统固碳能力的估算方法,今后的工作应注重海洋固碳整套观测技术、分析和估算方法研究,并建立海洋碳汇估算指标体系、指标标准体系、以及评价标准体系,为我国的碳"减排"、"增汇"国家需求提供技术支持。     

Estimation of phytoplankton production from space: current status and future potential of satellite remote sensing

A new generation of ocean colour satellites is now operational, with frequent observation of the global ocean. This paper reviews the potential to estimate marine primary production from satellite images. The procedures involved in retrieving estimates of phytoplankton biomass, as pigment concentrations, are discussed. Algorithms are applied to SeaWiFS ocean colour data to indicate seasonal variations in phytoplankton biomass in the Celtic Sea, on the continental shelf to the south west of the UK. Algorithms to estimate primary production rates from chlorophyll concentration are compared and the advantages and disadvantage discussed. The simplest algorithms utilise correlations between chlorophyll concentration and production rate and one equation is used to estimate daily primary production rates for the western English Channel and Celtic Sea; these estimates compare favourably with published values. Primary production for the central Celtic Sea in the period April to September inclusive is estimated from SeaWiFS data to be 102 gC m⁻² in 1998 and 93 gC m⁻² in 1999; published estimates, based on in situ incubations, are ca. 80 gC m⁻². The satellite data demonstrate large variations in primary production between 1998 and 1999, with a significant increase in late summer in 1998 which did not occur in 1999. Errors are quantified for the estimation of primary production from simple algorithms based on satellite-derived chlorophyll concentration. These data show the potential to obtain better estimates of marine primary production than are possible with ship-based methods, with the ability to detect short-lived phytoplankton blooms. In addition, the potential to estimate new production from satellite data is discussed.