海洋浮游植物铁限制的研究进展

介绍了海洋铁限制的研究进展和它对全球气候的影响,铁对海洋浮游植物的生长有着重要的作用,在高营养盐低叶绿素（HNLC）区域,铁是藻类生长的限制因子,海洋施铁对降低温室效应有着很好的前景,铁也是近岸赤潮发生的激发因子之一。     

铁对浮游植物生长影响的研究进展

浮游植物初级生产是海洋食物链的第一环节。光、温度和营养盐是影响海洋浮游植物光合作用的重要环境因子。这些环境因子的影响效应表现在藻类生长与繁殖的快慢。因此,人们对抑制浮游植物生长的营养盐是哪一种元素,众说纷坛。随着时间流逝,科学的发展趋势和研究结果使人们目前逐渐了解营养盐对浮游植物生长影响的机理和过程,同时,也了解营养盐生物地球化学过程。国内外学者对营养盐的研究也不断加深,研究结果也日新月异。本文主要阐述铁对浮游植物的增长影响和目前的研究进展。１　铁是浮游植物生长的限制因子的起源１９８２年,Ｇｏｒ…     

浮游植物生长上行效应的研究进展

归纳了海洋浮游植物生长营养盐限制的研究方法,综述了国内外关于海水中氮、磷、硅、铁等营养元素限制浮游植物生长的研究结果。在氮、磷营养元素限制时,海水中上述两个元素的浓度不仅限制了浮游植物的生长,而且还改变了浮游植物的种群结构,随着营养盐浓度的降低,浮游植物从小型向微型、微微型转变;而硅的缺乏,使浮游植物由硅藻向非硅藻转变;在大洋海域,铁的限制,甚至还影响到浮游植物对常量营养盐的吸收。铁施肥虽然促进了浮游植物的生长,降低了大气中CO2的浓度,缓解了温室效应,但同时也伴随着有毒藻类的生长以及DM S气体增加等带来的诸多问题。     

营养盐限制的唯一性因子探究

１营养盐限制因子的热点海洋中的几种元素 ,最引人关注的营养盐Ｎ ,Ｐ ,Ｓｉ和Ｆｅ对浮游植物生长是非常重要的 ,就象陆地上给植物施肥一样 ,营养盐限制浮游植物光合作用 ,在一些海域加了Ｎ ,Ｐ ,Ｓｉ,Ｆｅ,浮游植物生长就会旺盛、迅速。因此 ,这些营养盐对海域中浮游植物的生长起着重要的限制作用。营养盐Ｎ ,Ｐ ,Ｓｉ,Ｆｅ从海上的大气层、海底的上升流以及从陆地的河流中提供 ,从而决定了在海洋表面新的有机物质生产的速率 ［８］。这样 ,希望减少大气的ＣＯ ２,促进浮游植物的生长。增加限制营养盐的更大有用性是提高了海洋表面的初级生…     

铁、氮、磷、维生素B1和B12对海洋原甲藻的生长效应

４因子，２水平（铁为４水平）正交试验表明：络合铁和硝态氮对海洋原甲藻的生长有极显著的促进作用（Ｐ＜０．０１），维生不Ｂ１，Ｂ１２混合液有显著促进作用（Ｐ＜０．０５）。络合铁和硝态氮为大鹏湾浮游植物生长的营养限制因了。而磷酸盐的作用不显著。     

海水中铁的来源形态及其与浮游植物的相互关系

铁是浮游植物生长所必须的微量营养元素。铁在浮游植物对氮的吸收、固氮菌对N2的固定、叶绿素的合成、卟啉生物合成、光合作用电子的传输等生物过程发挥着非常重要的作用。浮游植物的光合作用固定了占全球近40％的碳，因此研究海洋中的铁的来源、形态及其与浮游植物的关系具有重要意义。本文论述了海水中铁的来源、形态及不同形态铁的相互转化，以及浮游植物需铁的生物过程、吸铁的方式、缺铁的生理生化指示和铁对浮游植物生长及群落组成的影响。     

海洋蓝藻固氮基因

氮是海洋浮游植物必需的大量元素，又是海洋浮游植物生长的主要限制因子之一，研究氮元素的补充机制对了解海洋生态系统有重要意义。海区氮营养盐的补充主要通过新生氮实现，其主要补充机制为径流输入、大气干湿沉降、水体混合、上升流及生物固氮作用。生物固氮是指固氮生物将大气中的氮气还原成氨的过程，对地球上氮循环和氮平衡具有不可替代的作用。海洋生物固氮作用是维持海洋初级生产力和新生产力的一个重要生态反应。海洋中固氮生物种类较多，主要包括固氮蓝藻、异养细菌和光合细菌。     

胶州湾营养盐限制浮游植物生长的初步模拟现场实验研究

于1998年8月至1999年2月，有胶州湾采用现场添加营养盐的实验方法初步研究了硝酸盐，磷酸盐，铁，硅酸盐对浮游植物生长的限制作用。分级叶绿素a的结果显示，在1998年夏季，氮（N）或单独，或与磷（P）同时限制浮游植物的生长；在1998的秋季则可能是N与P共同对浮游植物群落产生调控作用，而在1999年冬季，硅（Si)则是首要的限制浮游植物生长的营养元素。     

海水中溶解态铁的伏安极谱法与液芯波导光纤法测定

尽管铁为地壳中丰度位居第4的元素,但它在海水中的浓度却很低,浓度一般为0.05~2.0 nmd/dm3[1]。近年来,国内外的海洋科学家在进行海洋生态系统初级生产力、碳循环等重要生态过程及其对全球气候变化影响的研究中发现,海洋中的铁对浮游植物的生长、初级生产力的水平有着十分重要的影响甚至成为了“限制性因子”,在多个“高营养盐、低叶绿素(HNLC)”海区所实施的富铁实验的结果很好地印证了铁元素在海洋生态系统中的重要作用[2]。最新的研究成果表明近岸海水中的铁可能对浮游植物的种类组成起到决定性的作用,进而对包括氮、磷、硅在内的其他元素的生物地球化学行为产生深远的影响[3]。     

南沙群岛海域营养盐分布的研究

海洋中的营养盐是海洋浮游植物生长所必需的物质基础。营养盐、光照和温度是初级生产力的主要影响因素。海水中不同的营养组成和浓度比，不但会影响生物生产力，同时会对浮游植物的群落结构产生调节作用，影响着海洋生态系统的结构。因此，作为了解海洋生态系统生产过程的关键过程，海洋中的营养盐状况和循环研究，越来越得到生态学家的重视。诸多的研究表明，海洋中影响浮游植物生长的限制因子一般为Ｎ或Ｐ ,在中国沿海则主要是Ｐ的限制［１］。南沙群岛海域位于南海的南部，作为典型的热带海区，其营养盐状况与我国的温带地区是不同的。作…     

Growth stimulation and inhibition of natural phytoplankton communities by model organic ligands in the western subarctic Pacific

The influence of organic ligands on natural phytoplankton growth was investigated in high-nitrate low-chlorophyll (HNLC) waters and during a phytoplankton bloom induced by a mesoscale iron enrichment experiment (SEEDS II) in the western subarctic Pacific. The growth responses of the phytoplankton in the treatments with iron complexed with model ligand were compared with those with inorganic iron or a control. Desferrioxamine B and protoporphyrin IX were used as models for hydroxamate-type siderophore and tetrapyrrole-type cell breakdown ligand, respectively. In the HNLC water, iron associated with protoporphyrin IX especially stimulated smaller phytoplankton (<10 μm) growth, 1.5-fold more than did inorganic iron. Surprisingly, only the addition of protoporphyrin IX stimulated small phytoplankton growth, suggesting that these cell breakdown ligands might be more bioavailable for them. The protoporphyrin IX’s stimulatory effect on small phytoplankton was not observed during bloom decline phase. The growth of phytoplankton was inhibited in the treatment with desferrioxamine B-complexed iron, suggesting its low bioavailability for the natural phytoplankton community. Its inhibitory effects were particularly pronounced in pico-eukaryotic phytoplankton. During the iron-induced bloom, the phytoplankton’s iron-stress response gradually increased with the desferrioxamine B concentration, suggesting that the competition for iron complexation between natural ligands and desferrioxamine B affected phytoplankton growth. However, the pico-eukaryotes did seem better able to utilize the desferrioxamine B-complexed iron during the bloom-developing phase. These results indicate that the iron bioavailability for phytoplankton differs between bloom-developing and bloom-decline phases.     

Persistence of iron limitation in the western subarctic Pacific SEEDS II mesoscale fertilization experiment

The cumulative evidence from more than a dozen mesoscale iron-enrichment studies in high nitrate low chlorophyll (HNLC) waters demonstrates that iron limitation is widespread and very likely affects atmospheric carbon dioxide and thus global climate. However, the responses of microphytoplankton (>20 μm), predominantly diatoms, vary greatly among these mesoscale experiments even though similar amounts of iron were added, making it difficult to quantitatively incorporate iron effects into global climate models. Nowhere is this difference more dramatic than between the massive bloom observed during Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study (SEEDS) I and the order of magnitude smaller ecosystem response in SEEDS II; two mesocale experiments performed in the same HNLC region of the western subarctic Pacific in different years. Deckboard incubation experiments initiated during the early, middle, and late stages of the 32-day SEEDS II experiment show that while the two iron infusions increased phytoplankton growth, diatoms remained significantly limited by iron availability, despite total dissolved Fe concentrations in the patch being well above the diffusion-limited threshold for rapid diatom growth. This iron limitation was apparent <6 days after the initial iron infusion and was not alleviated by the second, smaller iron infusion. In contrast, smaller phytoplankton (<20 μm) showed a more restricted response to further iron amendments, indicating that their iron nutrition was near optimal. Iron complexed to desferrioximine B, a commonly available siderophore produced by at least one marine bacterium, was poorly available to diatoms throughout the patch evolution, indicating that these diatoms lacked the ability to induce high-affinity iron uptake systems. These results suggest that the strong organic complexation of Fe(III) observed in the SEEDS II-fertilized patch was not compatible with rapid diatom growth. In contrast, iron associated with protoporphyrin IX, a weaker iron complexing ligand of a class hypothesized to be representative of recycled iron species, was readily available to diatoms. Our findings demonstrate that a persistence of iron limitation was the primary factor underlying the comparatively small diatom response during SEEDS II. This continued growth limitation would have increased the importance of mesozooplankton grazing as a controlling factor in the SEEDS II ecosystem response.     

Physical limits on iron uptake mediated by siderophores or surface reductases

The excretion of siderophores and the reduction of organic iron-complexes at the cell surface are common reactions of terrestrial plants, fungi and bacteria in response to low availability of iron. However, there is much less evidence for the use of these strategies by marine phytoplankton. It has been argued that siderophore excretion is inefficient in an aquatic environment due to rapid diffusion. This study examines how diffusion and chemical reactions in the microenvironment of a phytoplankton cell influence the efficiency of both strategies to increase the bioavailability of iron and to reduce iron stress. A numerical model of the cell surroundings is presented that calculates the concentration distribution for different iron species and allows to study the effect of siderophores or surface reductases. It calculates the efficiency of these mechanisms, defined as the quotient between the increase in iron uptake rate and the excretion rate of siderophores or electrons, needed to obtain this increase. The dependence of this efficiency on rates of iron coordination reactions, on diffusivity, and on the kinetics of iron uptake is discussed with the aid of some analytical calculations.     

一氧化氮和铁对海洋微藻生长的交互影响

在实验室进行了一氧化氮(NO)和铁对海洋微藻中肋骨条藻(Skeletonema costatum)生长影响实验.实验结果如下:在铁限制情况下,外源NO能明显提高微藻的生长速率,增加生物量,但NO和铁对浮游植物生长影响是一个复杂的过程,它受到NO的浓度、培养液的营养水平、铁的浓度等因素的影响,同时培养液中铁的浓度也直接影响到微藻的生长和NO的释放量.因此,一氧化氮与铁在浮游植物生长过程中是交互影响的.     

Iron nutritional status of the phytoplankton assemblage in the Okhotsk Sea during summer

To clarify the iron (Fe) nutritional status of the phytoplankton assemblage in the Okhotsk Sea, we conducted incubation experiments in summer 2006. Replicate surface seawater samples with the natural plankton community were incubated with three treatments: Fe enrichment; addition of the strong Fe chelator siderophore desferriferrioxime B (DFB) which strips Fe from the biologically accessible pool; and as a control, no addition. To prevent macronutrient limitation, we added surplus nutrients to all treatments. At all 4 stations in Sakhalin Bay near the mouth of the Amur River and around the east of Sakhalin Island, net specific growth rate showed no significant difference between the control and +Fe treatment, and was repressed in +DFB treatment both in large- and small-sized phytoplankton. These findings indicate that these waters contain sufficient bioavailable Fe and that the Amur River plume which is transported by the east Sakhalin current is a major source of the Fe. In the Bussol’ Strait, net specific growth rate in the control was significantly higher than +DFB treatment, suggesting a supply of bioavailable Fe through intense vertical mixing at this site. Iron enrichment treatment stimulated the net specific growth rate of large-sized phytoplankton, indicating that Fe still limits the growth for the large-sized phytoplankton assemblage, but not for small-sized phytoplankton, in this area. An index of Fe availability was defined to quantify the degree of ambient Fe availability in each station, and it revealed the spatial variability of ambient Fe availabilities among the sites.     

Mutual effects of nitric oxide and iron on the growth of marine algae

Experiments on the effects of nitric oxide （NO） and iron on the growth of marine microalgae Skeletonema costatum were conducted. The results are as follows： exogenous NO could increase the growth rate of marine algae and raise the biomass remarkably under iron-deficient conditions. But it was a complicated process that the phytoplankton growth was influenced by NO and iron, which was controlled by the NO concentration, the nutrition level of the culture medium and the iron concentration, etc. Meanwhile, the iron concentration in the medium also has a direct influence on the growth and NO release capacity of the algae. Therefore, the effects of NO and iron on the growth of marine phytoplankton were mutual.     

北太平洋大气沉降的时空特征及其对副极区海洋生态系统的影响

北太平洋副极地海区作为全球海洋三个高营养盐低叶绿素(high nutrient and low chlorophyll, HNLC)海区之一, 其浮游植物生长受到微量元素铁的限制。对于开阔大洋, 大气沉降是海洋表层铁的一个重要来源, 铁元素沉降进入海洋后能够促进浮游植物生长, 进而引起海洋初级生产力和生物泵的响应。本文利用SPRINTARS(Spectral Radiation-Transport Model for Aerosol Species)模式的时长为20a的日均大气沉降数据, 对北太平洋海区大气沉降的时空特征进行了分析。结果表明, 进入北太平洋海区的大气沉降量为26.81Tg·a^-1, 并且存在显著的季节变化: 春季最高, 冬季最低, 5月份进入海洋的沉降量达到峰值。大气沉降主要来源于陆地区域, 在风场的驱动下向海洋传输, 因此大气沉降量的空间分布呈现出西高东低的特征。本文以2010年8月中旬卫星观测到的一次强沙尘(即高大气沉降量)事件为例, 研究了大气沙尘的传播路径。进一步结合2001年4月9—12日及2008年4月20—22日的沙尘事件, 分析了西北太平洋K2站位(47°N, 160°E)附近海域海洋初级生产力对大气沉降——沙尘事件的响应。结果表明, 三次沙尘事件后, K2站位的颗粒有机碳通量、叶绿素浓度均有明显增加, 即沙尘事件对北太平洋副极区海洋初级生产力存在促进作用。     

Development of a robust marine ecosystem model to predict the role of iron in biogeochemical cycles: A comparison of results for iron-replete and iron-limited areas,and the SOIREE iron-enrichment experiment

A new mixed layer multi-nutrient ecosystem model, incorporating diatoms, non-diatoms and zooplankton, is described that models the role of iron in marine biogeochemical cycles. The internal cell biochemistry of the phytoplankton is modelled using the mechanistic model of Flynn [2001. A mechanistic model for describing dynamic multi-nutrient, light, temperature interactions in phytoplankton. Journal of Plankton Research 23, 977–997] in which the internal cell concentrations of chlorophyll, nitrogen, silica, and iron are all dynamic variables that respond to external nutrient concentrations and light levels. Iron stress in phytoplankton feeds back into chlorophyll synthesis and changes in photosynthetic unit (PSU) size, thereby reducing their growth rate. Because diatom silicon metabolism is inextricably linked with cell division, diatom population density (cell m⁻³) is modelled as well as C biomass. An optimisation technique was used to fit the model to three time-series datasets at Biotrans (47°N, 20°W) and Kerfix (50°40′S, 68°25′E) and the observations for the Southern Ocean Iron-Release Experiment (SOIREE) iron-enrichment experiment (61°S, 140°E). The model gives realistic simulations of the annual cycles of nutrients, phytoplankton, and primary production at Biotrans and Kerfix and can also accurately simulate an iron fertilisation experiment. Specifically, the model predicts the high values of diatom Si:N and Si:C ratios observed in areas where iron is a limiting factor on algal growth. In addition, the model results at Kerfix confirm previous suggestions that underwater light levels have a more limiting effect on phytoplankton growth than iron supply. The model is also used to calculate C budgets and C and Si export from the mixed layer. The implications of these results for developing biogeochemical models incorporating the role of iron are discussed.     

A process-oriented model study of equatorial Pacific phytoplankton: The role of iron supply and tropical instability waves

The response of phytoplankton growth to iron supply and its modulation by large-scale circulation and tropical instability waves (TIWs) in the eastern equatorial Pacific has been investigated with an ocean biogeochemical model. This process study shows that iron can be efficiently advected from the New Guinea shelf through the Equatorial Undercurrent (EUC) to the eastern Pacific. The presence of a continental iron source is necessary for the maintenance of the observed subsurface iron maximum in the EUC core. In the eastern Pacific region, phytoplankton production is enhanced when additional iron is available in the EUC. Simulated phytoplankton variability is linked to TIWs activity, as revealed by a wavelet analysis of the total autotrophic carbon. The net local effect of the waves on phytoplankton can be either positive or negative depending on several factors. When the iron nutricline is sufficiently shallow to be reached by the wave vertical scale, the effect of the waves is to enhance iron availability in the euphotic zone leading to a net local increase of phytoplankton biomass. We therefore suggest that the local maxima of phytoplankton observed in moorings off the Equator in the eastern Pacific might be not only the result of concentration mechanisms, but also the result of an increase in local production sustained by advected iron.     

长江口理化因子影响初级生产力的探索Ⅰ.营养盐限制的判断方法和法则在长江口水域应用

营养盐限制浮游植物的生长是近年来国际研究的热点。探讨了目前营养盐限制的判断方法，认为仅靠氮和磷比值来得到磷限制或氮限制的结论是不完善的。根据营养盐限制的判断方法和法则，对长江口水域进行综合分析，认为在长江口及其附近海域，磷不是浮游植物生长的限制因子，而且氮、磷的浓度值都高于限制浮游植物生长的阈值，满足浮游植物的生长。通过1985年8月至1986年8月的长江口调查数据分析，发现从长江河口到远海水域的有些断面上，磷酸盐浓度并不一定离岸越远越低，且也没有周期性的季节变化。因此，尚须对河口区磷酸盐来源作进一步研究。