Reduced nitrogen fixation in the glacial ocean inferred from changes in marine nitrogen and phosphorus inventories.

To explain the lower atmospheric CO2 concentrations during glacial periods, it has been suggested that the productivity of marine phytoplankton was stimulated by an increased flux of iron-bearing dust to the oceans. One component of this theory is that iron-an essential element/nutrient for nitrogen-fixing organisms-will increase the rate of marine nitrogen fixation, fuelling the growth of other marine phytoplankton and increasing CO2 uptake. Here we present data that questions this hypothesis. From a sediment core off the northwestern continental margin of Mexico, we show that denitrification and phosphorite formation-processes that occur in oxygen-deficient upwelling regions, removing respectively nitrogen and phosphorus from the ocean-declined in glacial periods, thus increasing marine inventories of nitrogen and phosphorus. But increases in phosphorus were smaller and less rapid, leading to increased N/P ratios in the oceans. Acknowledging that phytoplankton require nitrogen and phosphorus in constant proportions, the Redfield ratio, and that N/P ratios greater than the Redfield ratio are likely to suppress nitrogen fixation, we suggest therefore that marine productivity did not increase in glacial periods in response to either increased nutrient inventories or greater iron supply.     

Diatom carbon export enhanced by silicate upwelling in the northeast Atlantic

Diatoms are unicellular or chain-forming phytoplankton that use silicon (Si) in cell wall construction. Their survival during periods of apparent nutrient exhaustion enhances carbon sequestration in frontal regions of the northern North Atlantic. These regions may therefore have a more important role in the 'biological pump' than they have previously been attributed, but how this is achieved is unknown. Diatom growth depends on silicate availability, in addition to nitrate and phosphate, but northern Atlantic waters are richer in nitrate than silicate. Following the spring stratification, diatoms are the first phytoplankton to bloom. Once silicate is exhausted, diatom blooms subside in a major export event. Here we show that, with nitrate still available for new production, the diatom bloom is prolonged where there is a periodic supply of new silicate: specifically, diatoms thrive by 'mining' deep-water silicate brought to the surface by an unstable ocean front. The mechanism we present here is not limited to silicate fertilization; similar mechanisms could support nitrate-, phosphate- or iron-limited frontal regions in oceans elsewhere.     

Spatial coupling of nitrogen inputs and losses in the ocean

Nitrogen fixation is crucial for maintaining biological productivity in the oceans, because it replaces the biologically available nitrogen that is lost through denitrification. But, owing to its temporal and spatial variability, the global distribution of marine nitrogen fixation is difficult to determine from direct shipboard measurements. This uncertainty limits our understanding of the factors that influence nitrogen fixation, which may include iron, nitrogen-to-phosphorus ratios, and physical conditions such as temperature. Here we determine nitrogen fixation rates in the world's oceans through their impact on nitrate and phosphate concentrations in surface waters, using an ocean circulation model. Our results indicate that nitrogen fixation rates are highest in the Pacific Ocean, where water column denitrification rates are high but the rate of atmospheric iron deposition is low. We conclude that oceanic nitrogen fixation is closely tied to the generation of nitrogen-deficient waters in denitrification zones, supporting the view that nitrogen fixation stabilizes the oceanic inventory of fixed nitrogen over time.     

Effect of natural iron fertilization on carbon sequestration in the Southern Ocean

The availability of iron limits primary productivity and the associated uptake of carbon over large areas of the ocean. Iron thus plays an important role in the carbon cycle, and changes in its supply to the surface ocean may have had a significant effect on atmospheric carbon dioxide concentrations over glacial-interglacial cycles. To date, the role of iron in carbon cycling has largely been assessed using short-term iron-addition experiments. It is difficult, however, to reliably assess the magnitude of carbon export to the ocean interior using such methods, and the short observational periods preclude extrapolation of the results to longer timescales. Here we report observations of a phytoplankton bloom induced by natural iron fertilization--an approach that offers the opportunity to overcome some of the limitations of short-term experiments. We found that a large phytoplankton bloom over the Kerguelen plateau in the Southern Ocean was sustained by the supply of iron and major nutrients to surface waters from iron-rich deep water below. The efficiency of fertilization, defined as the ratio of the carbon export to the amount of iron supplied, was at least ten times higher than previous estimates from short-term blooms induced by iron-addition experiments. This result sheds new light on the effect of long-term fertilization by iron and macronutrients on carbon sequestration, suggesting that changes in iron supply from below--as invoked in some palaeoclimatic and future climate change scenarios--may have a more significant effect on atmospheric carbon dioxide concentrations than previously thought.     

Climate-driven trends in contemporary ocean productivity

Contributing roughly half of the biosphere's net primary production (NPP), photosynthesis by oceanic phytoplankton is a vital link in the cycling of carbon between living and inorganic stocks. Each day, more than a hundred million tons of carbon in the form of CO2 are fixed into organic material by these ubiquitous, microscopic plants of the upper ocean, and each day a similar amount of organic carbon is transferred into marine ecosystems by sinking and grazing. The distribution of phytoplankton biomass and NPP is defined by the availability of light and nutrients (nitrogen, phosphate, iron). These growth-limiting factors are in turn regulated by physical processes of ocean circulation, mixed-layer dynamics, upwelling, atmospheric dust deposition, and the solar cycle. Satellite measurements of ocean colour provide a means of quantifying ocean productivity on a global scale and linking its variability to environmental factors. Here we describe global ocean NPP changes detected from space over the past decade. The period is dominated by an initial increase in NPP of 1,930 teragrams of carbon a year (Tg C yr(-1)), followed by a prolonged decrease averaging 190 Tg C yr(-1). These trends are driven by changes occurring in the expansive stratified low-latitude oceans and are tightly coupled to coincident climate variability. This link between the physical environment and ocean biology functions through changes in upper-ocean temperature and stratification, which influence the availability of nutrients for phytoplankton growth. The observed reductions in ocean productivity during the recent post-1999 warming period provide insight on how future climate change can alter marine food webs.     

Iron and phosphorus co-limit nitrogen fixation in the eastern tropical North Atlantic

The role of iron in enhancing phytoplankton productivity in high nutrient, low chlorophyll oceanic regions was demonstrated first through iron-addition bioassay experiments and subsequently confirmed by large-scale iron fertilization experiments. Iron supply has been hypothesized to limit nitrogen fixation and hence oceanic primary productivity on geological timescales, providing an alternative to phosphorus as the ultimate limiting nutrient. Oceanographic observations have been interpreted both to confirm and refute this hypothesis, but direct experimental evidence is lacking. We conducted experiments to test this hypothesis during the Meteor 55 cruise to the tropical North Atlantic. This region is rich in diazotrophs and strongly impacted by Saharan dust input. Here we show that community primary productivity was nitrogen-limited, and that nitrogen fixation was co-limited by iron and phosphorus. Saharan dust addition stimulated nitrogen fixation, presumably by supplying both iron and phosphorus. Our results support the hypothesis that aeolian mineral dust deposition promotes nitrogen fixation in the eastern tropical North Atlantic.     

Controls on tropical Pacific Ocean productivity revealed through nutrient stress diagnostics

In situ enrichment experiments have shown that the growth of bloom-forming diatoms in the major high-nitrate low-chlorophyll (HNLC) regions of the world's oceans is limited by the availability of iron. Yet even the largest of these manipulative experiments represents only a small fraction of an ocean basin, and the responses observed are strongly influenced by the proliferation of rare species rather than the growth of naturally dominant populations. Here we link unique fluorescence attributes of phytoplankton to specific physiological responses to nutrient stress, and use these relationships to evaluate the factors that constrain phytoplankton growth in the tropical Pacific Ocean on an unprecedented spatial scale. On the basis of fluorescence measurements taken over 12 years, we delineate three major ecophysiological regimes in this region. We find that iron has a key function in regulating phytoplankton growth in both HNLC and oligotrophic waters near the Equator and further south, whereas nitrogen and zooplankton grazing are the primary factors that regulate biomass production in the north. Application of our findings to the interpretation of satellite chlorophyll fields shows that productivity in the tropical Pacific basin may be 1.2-2.5 Pg C yr(-1) lower than previous estimates have suggested, a difference that is comparable to the global change in ocean production that accompanied the largest El Ni?o to La Ni?a transition on record.     

坡面氮、磷流失特征分析及预测

利用室内人工降雨系统和大型可变坡土槽,模拟了裸土坡地在不同的初始土壤营养物质量分数、径流量、坡度及植被覆盖率下的产污过程,分析了典型降雨过程中氮、磷质量浓度随时间的变化过程以及径流中营养物质的流失规律.结果表明:径流中氮、磷质量浓度受表层土壤营养物的质量分数影响很大;氮、磷质量浓度在产流初期迅速下降,随后下降速度趋于平稳;在多数情况下,径流中硝酸盐氮的质量浓度高于氨氮质量浓度,亚硝酸盐氮的质量浓度很低.在物理机理研究的基础上提出了氮、磷的污染负荷模型.     

Reduced mixing generates oscillations and chaos in the oceanic deep chlorophyll maximum

Deep chlorophyll maxima (DCMs) are widespread in large parts of the world's oceans. These deep layers of high chlorophyll concentration reflect a compromise of phytoplankton growth exposed to two opposing resource gradients: light supplied from above and nutrients supplied from below. It is often argued that DCMs are stable features. Here we show, however, that reduced vertical mixing can generate oscillations and chaos in phytoplankton biomass and species composition of DCMs. These fluctuations are caused by a difference in the timescales of two processes: (1) rapid export of sinking plankton, withdrawing nutrients from the euphotic zone and (2) a slow upward flux of nutrients fuelling new phytoplankton production. Climate models predict that global warming will reduce vertical mixing in the oceans. Our model indicates that reduced mixing will generate more variability in DCMs, thereby enhancing variability in oceanic primary production and in carbon export into the ocean interior.     

Pumping of nutrients to ocean surface waters by the action of propagating planetary waves

Primary productivity in the oceans is limited by the lack of nutrients in surface waters. These nutrients are mostly supplied from nutrient-rich subsurface waters through upwelling and vertical mixing, but in the ocean gyres these mechanisms do not fully account for the observed productivity. Recently, the upward pumping of nutrients, through the action of eddies, has been shown to account for the remainder of the primary productivity; however, these were regional studies which focused on mesoscale (100-km-scale) eddies. Here we analyse remotely sensed chlorophyll and sea-surface-height data collected over two years and show that 1,000-km-scale planetary waves, which propagate in a westward direction in the oceans, are associated with about 5 to 20% of the observed variability in chlorophyll concentration (after low-frequency and large-scale variations are removed from the data). Enhanced primary production is the likely explanation for this observation, and if that is the case, propagating disturbances introduce nutrients to surface waters on a global scale--similar to the nutrient pumping that occurs within distinct eddies.     

Biomarker reconstruction of phytoplankton productivity and community structure changes in the middle Okinawa Trough during the last 15 ka

Biomarkers have been widely used to reconstruct phytoplankton productivity and community structure changes, and this method has been applied for the first time in the middle Okinawa Trough during the transition from the last deglaciation to the Holocene. The total content of all marine phytoplankton biomarkers, used as a total productivity indicator, reveals higher productivity during the deglaciation. The ratios of the biomarkers are used as community structure indicators which show that, compared with the Holocene, the contribution from haptophytes decreased while the contributions from diatoms and dinoflagellates increased during the deglaciation. The increased productivity during the deglaciation was likely caused by the stronger winter monsoon. Also increased nutrient supply from terrestrial sources contributes to the higher productivity due to lower sea-level, which is consistent with higher terrestrial biomarker （long-chain n-alkanols） content. These changes in the nutrient supply also con- tributed to the community structure changes in the Okinawa Trough.     

东、黄海典型海区浮游植物对营养盐添加的响应

2002年9月,通过现场营养盐添加实验研究了长江口外海(E4测站)、东海陆架中部海区(E6测站)以及黄海冷水团(E2测站)分粒级叶绿素a对营养盐添加的响应.结果显示:E6测站加磷与加氮磷组均在120h出现了叶绿素峰值响应,表明9月该站同时受氮磷限制.E4测站加磷组在第1天培养结束时就出现了叶绿素响应,证实了磷限制的存在.E2测站由于培养时间所限,加氮与加磷组均未观测到叶绿素明显的峰值响应.不同海区浮游植物对响应时间不尽相同.营养盐的添加在一定程度上改变了浮游植物的粒级结构.     

Global biodiversity patterns of marine phytoplankton and zooplankton

Although the oceans cover 70% of the Earth's surface, our knowledge of biodiversity patterns in marine phytoplankton and zooplankton is very limited compared to that of the biodiversity of plants and herbivores in the terrestrial world. Here, we present biodiversity data for marine plankton assemblages from different areas of the world ocean. Similar to terrestrial vegetation, marine phytoplankton diversity is a unimodal function of phytoplankton biomass, with maximum diversity at intermediate levels of phytoplankton biomass and minimum diversity during massive blooms. Contrary to expectation, we did not find a relation between phytoplankton diversity and zooplankton diversity. Zooplankton diversity is a unimodal function of zooplankton biomass. Most strikingly, these marine biodiversity patterns show a worldwide consistency, despite obvious differences in environmental conditions of the various oceanographic regions. These findings may serve as a new benchmark in the search for global biodiversity patterns of plants and herbivores.     

Rapid and early export of Phaeocystis antarctica blooms in the Ross Sea, Antarctica

The Southern Ocean is very important for the potential sequestration of carbon dioxide in the oceans and is expected to be vulnerable to changes in carbon export forced by anthropogenic climate warming. Annual phytoplankton blooms in seasonal ice zones are highly productive and are thought to contribute significantly to pCO2 drawdown in the Southern Ocean. Diatoms are assumed to be the most important phytoplankton class with respect to export production in the Southern Ocean; however, the colonial prymnesiophyte Phaeocystis antarctica regularly forms huge blooms in seasonal ice zones and coastal Antarctic waters. There is little evidence regarding the fate of carbon produced by P. antarctica in the Southern Ocean, although remineralization in the upper water column has been proposed to be the main pathway in polar waters. Here we present evidence for early and rapid carbon export from P. antarctica blooms to deep water and sediments in the Ross Sea. Carbon sequestration from P. antarctica blooms may influence the carbon cycle in the Southern Ocean, especially if projected climatic changes lead to an alteration in the structure of the phytoplankton community.     

Lake eutrophication and its ecosystem response

China is a country with many lakes, about one-third of which are freshwater mainly distributed in the middle and lower reaches of the Yangtze River. Currently most of the lakes are mesotrophic or eutrophic. Lake eutrophication has become one of the major ecological and environmental problems faced by lakes in China and can lead to a series of abnormal ecosystem responses, including extinction of submerged plants, frequent occurrence of cyanobacterial blooms, increased microbial biomass and productivity, decreased biodiversity, accelerated cycles, and a change in the efficient use of nutrients. With development of eutrophication, the whole lake ecosystem suffers decreased biodiversity, simplification of biotic community structure, instability of the ecosystem, and ultimately the clear-water, macrophyte-dominated ecosystem gradually shifts to a turbid-water, algae-dominated ecosystem. This ecosystem succession mechanism is speculated to be caused by different nutrient utilization efficiencies of macrophytes and phytoplankton. The ultimate ecosystem succession trend of seriously eutrophic lakes is that a phytoplankton-dominated autotrophic lake shifts to a heterotrophic lake dominated by micro-organisms, protozoans.     

营养盐传感器在海洋监测中的研究进展

营养盐是海洋浮游植物生长所必需的物质基础。测定海水中营养盐的分布对于理解地球化学循环,预测藻华爆发等环境灾害有重要的意义。营养盐在线传感器在过去的几十年中取得了较大的进展。本文综述了营养盐传感器在海水监测中的研究进展,介绍和比较了不同营养盐传感器的检测原理、范围和检测限,并讨论了营养盐传感器的技术瓶颈和发展趋势。     

Nitrogen loss from unpolluted South American forests mainly via dissolved organic compounds.

Conceptual and numerical models of nitrogen cycling in temperate forests assume that nitrogen is lost from these ecosystems predominantly by way of inorganic forms, such as nitrate and ammonium ions. Of these, nitrate is thought to be particularly mobile, being responsible for nitrogen loss to deep soil and stream waters. But human activities-such as fossil fuel combustion, fertilizer production and land-use change-have substantially altered the nitrogen cycle over large regions, making it difficult to separate natural aspects of nitrogen cycling from those induced by human perturbations. Here we report stream chemistry data from 100 unpolluted primary forests in temperate South America. Although the sites exhibit a broad range of environmental factors that influence ecosystem nutrient cycles (such as climate, parent material, time of ecosystem development, topography and biotic diversity), we observed a remarkably consistent pattern of nitrogen loss across all forests. In contrast to findings from forests in polluted regions, streamwater nitrate concentrations are exceedingly low, such that nitrate to ammonium ratios were less than unity, and dissolved organic nitrogen is responsible for the majority of nitrogen losses from these forests. We therefore suggest that organic nitrogen losses should be considered in models of forest nutrient cycling, which could help to explain observations of nutrient limitation in temperate forest ecosystems.     

Endangered plants persist under phosphorus limitation

Nitrogen enrichment is widely thought to be responsible for the loss of plant species from temperate terrestrial ecosystems. This view is based on field surveys and controlled experiments showing that species richness correlates negatively with high productivity and nitrogen enrichment. However, as the type of nutrient limitation has never been examined on a large geographical scale the causality of these relationships is uncertain. We investigated species richness in herbaceous terrestrial ecosystems, sampled along a transect through temperate Eurasia that represented a gradient of declining levels of atmospheric nitrogen deposition--from approximately 50 kg ha(-1) yr(-1) in western Europe to natural background values of less than 5 kg ha(-1) yr(-1) in Siberia. Here we show that many more endangered plant species persist under phosphorus-limited than under nitrogen-limited conditions, and we conclude that enhanced phosphorus is more likely to be the cause of species loss than nitrogen enrichment. Our results highlight the need for a better understanding of the mechanisms of phosphorus enrichment, and for a stronger focus on conservation management to reduce phosphorus availability.     

香溪河库湾春季水华纵向分布对水层结构的响应

为研究三峡水库支流库湾春季水华暴发空间差异及成因,本研究于2010年4月18日对三峡水库干流和香溪河库湾的水动力、营养盐、浮游植物等因子进行了现场监测.结果显示：三峡水库支流春季水华浮游植物生物量及群落结构空间差异较大,库湾中上游暴发了严重的硅藻水华,而下游及三峡水库干流没有水华;库湾中复杂的分层异重流对水温结构产生显著影响,使其呈独特的垂向分层;水温分层纵向差异导致的光混比（Zeu/Zmix）纵向分布差异是引起水华情势和浮游植物群落结构纵向差异的主要原因.研究结果将为三峡水库支流水华机理和防控措施研究提供支撑.     

Consumer versus resource control of species diversity and ecosystem functioning

A key question in ecology is which factors control species diversity in a community. Two largely separate groups of ecologists have emphasized the importance of productivity or resource supply, and consumers or physical disturbance, respectively. These variables show unimodal relationships with diversity when manipulated in isolation. Recent multivariate models, however, predict that these factors interact, such that the disturbance diversity relationship depends on productivity, and vice versa. We tested these models in marine food webs, using field manipulations of nutrient resources and consumer pressure on rocky shores of contrasting productivity. Here we show that the effects of consumers and nutrients on diversity consistently depend on each other, and that the direction of their effects and peak diversity shift between sites of low and high productivity. Factorial meta-analysis of published experiments confirms these results across widely varying aquatic communities. Furthermore, our experiments demonstrate that these patterns extend to important ecosystem functions such as carbon storage and nitrogen retention. This suggests that human impacts on nutrient supply and food-web structure have strong and interdependent effects on species diversity and ecosystem functioning, and must therefore be managed together.