海藻营养代谢研究进展国家杰出青年科学——海藻营养代谢的调节

介绍影响海藻(尤其是大型海藻)主要营养盐(N,P,C)吸收的环境因子(如光照、温度及水流)对营养盐主动吸收过程中的能量提供，酶含量和活性的影响以及水流对藻体周围营养盐离子进入细胞的调节；化学因素(营养盐离子或分子的浓度、形式、海水介质的pH)对海藻选择吸收不同形式营养盐离子及其相互作用的调节过程；生物因素(藻体形态、组织的类型及海藻的年龄和营养史)通过细胞内不同水平营养库的积累和相互转变而对海藻营养吸收和同化的调节；N代谢与C代谢的生化偶连关系。     

环境因子对海洋藻类酶活性的影响及其应用

海洋藻类是海洋中的初级生产者,在其生长发育过程中体内各种各样的酶不断地催化着无数的生化反应,完成其新陈代谢过程,而决定这些酶促反应强弱的酶活性的高低会直接或间接地受到外界环境条件的影响.简要概述了物理因素(如光照、温度及水流)、化学因素(营养盐离子或分子的浓度、形式、海水介质的pH)、生物因素(藻体形态、组织的类型及海藻的年龄)对海洋藻类酶含量和活性的影响及其应用前景.     

环境因子对海洋藻类酶活性的影响及其应用

海洋藻类是海洋中的初级生产者,在其生长发育过程中体内各种各样的酶不断地催化着无数的生化反应,完成其新陈代谢过程,而决定这些酶促反应强弱的酶活性的高低会直接或间接地受到外界环境条件的影响.简要概述了物理因素(如光照、温度及水流)、化学因素(营养盐离子或分子的浓度、形式、海水介质的pH)、生物因素(藻体形态、组织的类型及海藻的年龄)对海洋藻类酶含量和活性的影响及其应用前景.     

藻体部位、藻质量、碳源、水流对铁钉菜和羊栖菜营养盐吸收的影响

采用实验室生态学研究方法,进行了藻体部位、藻质量、碳源质量浓度及水流流速对铁钉菜(Ishige foliaceaokamurai)和羊栖菜(Sargassum fusiforme)氮磷营养盐吸收影响的研究。结果表明:2种大型海藻不同部位对营养盐的吸收速率不同,铁钉菜下部的吸收速率最大,其次是固着器;羊栖菜顶部的吸收速率最大,其次是中部。铁钉菜藻质量1.0g的吸收速率最大,其次是1.5g;羊栖菜藻质量0.5 g的吸收速率最大,其次是1.0g;2种藻类2.0g和2.5g的吸收速率相差不多。碳源质量浓度为1200mg/L时,铁钉菜的吸收速率最大,其次是碳源800mg/L;碳源质量浓度为800mg/L时,羊栖菜的吸收速率最大,其次是碳源400mg/L;2种海藻对照组的吸收速率均相对较低。相同流速下,2种海藻均以PO4-P对照组的吸收速率最大;NO3-N、NH4-N吸收速率在流速为50cm/s时最大。     

不同配比饲料对刺参C、N、P营养盐收支的影响

本实验采用5种含有不同比例(即20∶80、40∶60、60∶40、80∶20和100∶0)的鼠尾藻和海泥的颗粒饲料喂养刺参(Apostichopus japonicus),并比较了不同饲料条件下刺参C、N、P营养盐的收支过程。结果表明:刺参C、N、P营养盐收支过程和饲料中鼠尾藻的含量显著相关(P<0.05)。刺参的摄食率和排粪率随着饲料中鼠尾藻含量的增加而显著降低(P<0.05);刺参的耗氧率、排氨率、排磷率,以及对C、N、P营养盐的摄入率、生长余力和吸收效率均随着饲料中鼠尾藻含量的增加而显著变大(P<0.05)。当饲料中鼠尾藻添加量大于60%时,刺参对营养元素的生长余力和吸收效率无显著变化(P>0.05),因此对于成参而言,饲料中鼠尾藻和海泥的比例为60∶40是较为理想的配制比例,既可以满足刺参生长代谢所需要的主要营养元素,又能够最大限度地减少刺参养殖活动营养盐的排放。     

浮游植物营养代谢研究新进展

本文概括介绍了浮游植物营养代谢研究的方法,并对其营养盐的吸收、同化、释放及其影响因素的研究做了简要的论述。     

陆源排污对邻近海域底栖海藻群落的影响

伴随着经济的发展,我国沿海海域富营养化日趋严重。为了明确富营养化对底栖海藻群落的影响及作用机制,本文选择了与大连凌水河口(污水河)毗邻的海藻床作为野外观测点,研究底栖海藻群落对营养盐自然衰减梯度的响应规律;在实验室内,选择了在排污口附近优势分布、营r-生态策略的绿藻缘管浒苔和仅在寡营养区域分布、营k-生态策略的红藻柔质仙菜作为实验材料,开展了营养吸收动力学和生长动力学研究。通过二者在营养盐吸收、利用和繁殖策略方面的比较研究,剖析了底栖海藻群落对不同营养环境的响应机制。结果显示:随着海水中营养盐浓度的降低,底栖海藻群落呈现种类数增加、优势种覆盖度降低的趋势。根据底栖海藻在群落水平对氮源营养的响应,认为现行海水水质标准中无机氮一类水质标准的限值应该由目前的14.29 μmol/L降低为6.69 μmol/L。在富营养环境中,营养盐浓度的上升促进了r-策略海藻幼体的竞争力和种群繁殖力,使其占据了大量的生态位,形成优势种群,导致底栖海藻多样性较低;在寡营养环境中,由于得不到充足的营养盐供应,r-策略海藻幼体的竞争力和种群繁殖力都受到制约,占据的空间生态位有限,为其他种类的生存提供了条件,而那些对营养盐需求较低但利用效率高的k-策略海藻则表现出更强的竞争力,在竞争中取得优势,能快速突破早期环境筛的限制,形成成体,因此,在寡营养海域,底栖海藻的多样性比较丰富。     

光照和营养盐胁迫对龙须菜生长及生化组成的影响

以龙须菜(Gracilaria lemaneiformis)为实验材料,分别研究了光照强度、营养盐胁迫对其生长及生化组成的影响。结果表明,在800～3000lx的范围内随光强的升高,龙须菜生长率增加,但藻红素(PE)、叶绿素a、蛋白质和可溶性总糖含量却降低,即低光强更利于龙须菜生化组分的积累。饥饿处理组、饱和处理组及正常对照组的生长率分别为0.9%,5.01%,3.06%。饥饿处理组的总C含量最高,总N及C/N比值高(>15),饱和处理组则相反,C/N比值<10,即存在C高N低、N高C低的现象。表明水体营养状况对龙须菜生长、体内生化组分有显著影响。总C、总N及C/N比都能很好地反映藻体的营养状态。饱和处理组藻体呈现深红色,饥饿处理组藻体色泽为浅黄色,因此也可根据藻体色泽的变化来估计藻体的营养状态。     

4种大型海藻单养和两两混养条件下生长以及营养盐的吸收利用

近年来海水富营养化导致近海海域大型海藻过量繁殖,为研究大型海藻与营养盐的相互关系,我们采用实验生态学的方法研究了4种大型海藻(孔石莼、小珊瑚藻、缘管浒苔、石花菜)单养和两两混养条件下的生长、营养盐(PO4-P和NO3-N)含量变化以及营养盐吸收利用情况。结果表明,单养条件下,孔石莼的湿重增长最多(46.4%),其相对增长率显著高于缘管浒苔、小珊瑚藻、石花菜。孔石莼+缘管浒苔、孔石莼+小珊瑚藻、孔石莼+石花菜混养体系中,孔石莼的相对增长率显著高于缘管浒苔、小珊瑚藻、石花菜,其他混养体系中两种大型海藻的相对增长率无显著差异。在单养条件下培养10d,海藻对PO4-P、NO3-N的利用率分别以孔石莼(85.4%)、缘管浒苔(72.6%)为最大,而石花菜对两种营养盐的利用率均最小(51.1%和57.1%)。在混养条件下,PO4-P、NO3-N的最高利用率分别出现在孔石莼+小珊瑚藻(86.3%)和缘管浒苔+小珊瑚藻(81.7%)体系中。无论单养还是混养,海藻对PO4-P和NO3-N的吸收速率都在第2d达到峰值。因此,大型海藻可以有效地去除营养盐,4种藻中,孔石莼和缘管浒苔分别是去除PO4-P和NO3-N的最佳大型海藻。本研究结果可以为利用大型海藻进行环境污染的治理提供支持。     

基于转录组测序对两种氮素营养条件下多形微眼藻氮代谢途径的解析

探究微藻的氮代谢通路对了解其对不同氮源利用的分子机制具有重要意义。本研究利用Illumina高通量测序技术对两种氮素营养条件下(硝酸氮和尿素)多形微眼藻的转录组进行分析,通过基因功能注释及数字基因表达谱分析,研究了多形微眼藻细胞内氮代谢的调控机制。结果检测出15种参与氮代谢的酶,对应76个编码基因,构建了多形微眼藻的氮代谢通路图。其中10个酶编码基因在两种不同氮素营养条件下存在差异表达,最显著的是谷氨酸合成酶、谷氨酸脱氢酶和谷氨酰胺合成酶相关基因。有机氮源(尿素)实验组中,多形微眼藻细胞内的硝酸盐还原酶、亚硝酸盐还原酶等基因的差异表达明显高于无机氮源(硝酸钠)实验组,表明当环境中的氮源为尿素时,会对多形微眼藻细胞内硝酸盐的转化和利用有一定影响。本研究初步阐述了硝酸盐、尿素的吸收转运对多形微眼藻细胞内氮代谢的影响机制,可为硅藻在不同氮素营养条件下的吸收利用机制及氮代谢响应研究提供依据。     

Formation and functional significance of storage products in cyanobacteria

The most common storage products of cyanobacteria are polyphosphate as a phosphorus storage compound, cyanophycin or phycobilin protein pigment as nitrogen storage products, and glycogen as a storage product of both carbon and energy. Nutrient uptake kinetics are regulated by the storage pools, and the patterns of regulation have a feedback effect on the amount of accumulated nutrient in the cells. Besides having a storage function the nutrient storage products are likely to act as metabolic sinks during conditions of energy stress. Regulation of storage products is especially strict in light‐limited cultures. By increasing the rate of polysaccharide formation during growth with short photoperiods, cyanobacteria are able to sustain relatively high growth rates. This effect is enhanced by keeping respiratory losses very low.     

Physiological acclimation by phytoplankton explains observed changes in Si and N uptake rates during the SERIES iron-enrichment experiment

During the SERIES iron-enrichment experiment in the eastern subarctic Pacific, after addition of iron and its subsequent depletion, the Si:N drawdown ratio increased at approximately the time that diatoms became iron limited. Laboratory studies have reported that this results from a decrease in the rate of N uptake together with a more moderate decrease in the rate of Si uptake for iron-limited cultures compared to iron-replete cultures. However, for SERIES Boyd et al. (Limnol. Oceanogr. 50 (2005)) reported an unexplained increase in the rate of Si uptake at the onset of iron limitation and suggested that studies of nutrient uptake kinetics should be undertaken in search of an explanation. We compare the classic Michealis–Menten (MM) kinetics to the recently developed optimal uptake (OU) kinetics (the SPONGE: Smith and Yamanaka. Limnol. Oceanogr. 52 (2007)) within a variable-composition model, which employs cell quotas for each relevant nutrient, applied to the multi-element (C, N, Si, Fe) dynamics during SERIES. Using the Monte Carlo Markov Chain, we fit two versions of the model (differing only in the equations for nutrient uptake) to the available data for nutrient concentrations, chlorophyll, biogenic silica and particulate organic carbon and specific growth rates.With either uptake kinetics, the model reproduces observed concentrations well for nutrients and somewhat less well for chlorophyll. The different uptake kinetics yield greater differences in modeled elemental composition of phytoplankton and biomass of phytoplankton and zooplankton, which are not directly constrained by data. MM kinetics cannot reproduce the observed increase in Si uptake rate as a function of the decreasing trend in concentration of silicic acid, and it predicts Si limitation throughout nearly all of the experiment after iron-fertilization. In contrast, OU kinetics reproduces the increase in Si uptake rate and matches the observation-based estimate for the timing of the return to iron limitation. The key assumption of the SPONGE, that uptake rates of all nutrients depend on physiological acclimation by phytoplankton as a function of the ambient concentration of the growth-limiting nutrient, was originally formulated for modeling chemostat experiments. We show that it also agrees with the observations from this field experiment and provides an explanation for the increases in Si uptake rate and Si:N drawdown ratio.     

Bottom-up control of phytoplankton growth in spring blooms in Central Yellow Sea,China

The source and significance of three nutrients – nitrogen, phosphorous and silicon – were investigated by a modified dilution method performed on seawater samples from the Central Yellow Sea (CYS), in spring blooming period of 2007. This modified dilution method accounted for the phytoplankton growth rate, microzooplankton grazing mortality rate, the internal and external nutrient pools, as well as nutrients supplied through remineralization by microzooplankton grazing. The results indicate that phytoplankton growth during the bloom is mostly contributed by internal nutrient pools (K_I=0.062–1.730). The external nutrient pools (K_E=<0–0.362) are also of importance for phytoplankton growth during the bloom at some sampling sites. Furthermore, the contribution of the recycled-nutrient pool by remineralization (K_R=<0–0.751) is significant when microzooplankton grazing rate was higher than 0.5 d⁻¹ during the spring phytoplankton blooms in the Central Yellow Sea. Compared with internal phosphorus, internal nitrogen and silicon contribute more to the phytoplankton production at most sampling stations.     

Preliminary study on different nutrient pools supplies for the phytoplankton growth in the Jiaozhou Bay in China in the fall of 2004

The source and significance of two nutrients, nitrogen and phosphorous, were investigated by a modified dilution method performed on seawater samples from the Jiaozhou Bay, in autumn 2004. This modified dilution method accounted for the phytoplankton growth rate, microzooplankton grazing mortality rate, the internal and external nutrient pools, as well as nutrient supplied through remineralization by microzooplankton. The results indicated that the phytoplankton net growth rate increased in turn from inside the bay, to outside the bay, to in the Xiaogang Harbor. The phytoplankton maximum growth rates and microzooplankton grazing mortality rates were 1.14 and 0.92 d-1 outside the bay, 0.42 and 0.32 d-1 inside the bay and 0.98 and 0.62 d-1 in the harbor respectively. Outside the bay, the remineralized nitrogen (Kr=24.49) had heavy influence on the growth of the phytoplankton. Inside the bay, the remineralized phosphorus(Kr=3.49) strongly affected the phytoplankton growth. In the harbor, the remineralized phosphorus (Kr=3.73) was in larger demand by phytoplankton growth. The results demonstrated that the different nutrients pools supplied for phytoplankton growth were greatly in accordance with the phytoplankton community structure, microzooplankton grazing mortality rates and environmental conditions. It is revealed that nutrient remineralization is much more important for the phytoplankton growth in the Jiaozhou Bay than previously believed.     

Phosphate Utilization in Phaeocystis pouchetii (Haptophyceae)

Abstract. .The phosphate uptake kinetics of single cells and colonies of the haptophycean alga Phaeocystis pouchetii were investigated in short- and long-term experiments. In phosphate-deprived cultures, the highest nutrient uptake rate (V_max) of both growth forms appeared to be similar. However, the half-saturation constant for phosphate uptake of the cells embedded in a colony (K_s= 3.08 μM) was considerably higher than that of the single cells (K_s= 0.31 μM). This is consistent with the existence of a diffusion barrier formed by the colony membrane and colonial mucus. Over a 24 h light/dark cycle, the phosphate assimilation rate in moderately limited colonies and single cells was nearly identical during the light period. In the following dark phase, colony cells maintained this uptake rate, whereas the uptake of the single cells diminished by 70 %. The high dark uptake rate in colonies ran parallel with the absorption and degradation of intracolonial carbon compounds, suggesting that the dissimilation of these compounds covers the energy requirements for phosphate uptake in the dark. In Phaeocystis cultures during a transition to phosphate depletion, a significant amount of phosphate was found in the colonial mucus, isolated by selective filtration. The possible absorption of phosphate by the mucus and the role of mucus in the energy budget seem to be an essential feature of phosphate uptake of colonies. Short-term measurement of phosphate uptake in colony cells may therefore underestimate the phosphate uptake of the whole colony.     

脉冲式营养盐输入对中肋骨条藻生长的影响

脉冲输入营养盐是陆源输入营养盐的一种方式。用室内模拟脉冲营养盐输入的方法,研究了脉冲营养盐输入对于典型赤潮藻中肋骨条藻(Skeletonema costatum)生长的影响。结果发现脉冲输入营养盐对于中肋骨条藻生长有明显的影响,营养盐脉冲输入的频率和中肋骨条藻生长波动的频率相同。每天输入一次营养盐中肋骨条藻出现藻密度峰值的时间要比每5天输入一次营养盐和每10天输入一次营养盐的中肋骨条藻要滞后,而且藻密度峰值也比后两种情况低。对于3种营养盐的吸收速率而言,每10天输入一次营养盐的中肋骨条藻的吸收速率最大,其次是每天输入一次营养盐的中肋骨条藻,最小的是每5天输入一次营养盐的中肋骨条藻;3种营养盐脉冲输入模式下,每5天输入一次营养盐的中肋骨条藻对于N盐和Si盐的营养需求最少。     

Vertical Nutrient Distributions in the Western North Pacific Ocean: Simple Model for Estimating Nutrient Upwelling, Export Flux and Consumption Rates

A one-dimensional, steady-state model has been developed to understand the factors controlling vertical distributions of nutrients such as nitrate and phosphate in the western North Pacific water columns. The model includes simple physics and some biogeochemical processes. Nutrients are supplied by upwelling of nutrient-rich deep waters with a constant upwelling velocity and nutrient regeneration due to decomposition of sinking particulate matter; the latter is expressed by an exponential-type export flux. Nutrients are consumed in the water column due to uptake by marine organisms, represented by a first-order substrate kinetics. The consumption rate constant is given as an exponential function of depth. The model has been applied to a data set of WOCE (World Ocean Circulation Experiment) P9 one-time measurements observed in the western North Pacific. The calculated curves fit well to observed vertical nutrient profiles from 100 m depth to over 2,500 m depth at 35 stations from 19°N to 33°30′ N along 137°E with correlation factors of greater than 0.998. A modified model, including a correction term representing a depth-dependent upwelling velocity, can reproduce observed vertical nutrient profiles at 32 stations from 5°N to 18°30′ N along 137°E with correlation factors greater than 0.993. The results support the hypothesis that most of the vertical nutrient profiles in the western North Pacific are controlled by particle export flux, consumption rate, remineralization rate and upwelling velocity. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inorganic nitrogen acquisition by the tropical seagrass Halophila stipulacea

The tropical seagrass Halophila stipulacea is dominant in most regions of the Indo‐Pacific and the Red Sea and was introduced into the Mediterranean Sea after the opening of the Suez canal. The species is considered invasive in the Mediterranean Sea and has been progressively colonizing new areas westward. Growth and photosynthetic responses of H. stipulacea have been described but no information is yet available on the nitrogen nutrition of the species. Here we simultaneously investigated the uptake kinetics of ammonium and nitrate and the internal translocation of incorporated nitrogen in H. stipulacea using ¹⁵N‐labelled substrates across a range of N_i levels (5, 25, 50 and 100 μm ). The ammonium uptake rates exceeded the nitrate uptake rates 100‐fold, revealing a limited capacity of H. stipulacea to use nitrate as an alternative nitrogen source. The uptake rates of ammonium by leaves and roots were comparable up to 100 μm ¹⁵NH₄Cl. At this concentration, the leaf uptake rate was 1.4‐fold higher (6.22 ± 0.70 μmol·g^?1 DW h^?1) than the root uptake rate (4.54 ± 0.28 μmol·g^?1 DW h^?1). The uptake of ammonium followed Michaelis–Menten kinetics, whereas nitrate uptake rates were relatively constant at all nutrient concentrations. The maximum ammonium uptake rate (V_max) and the half‐saturation constant (K_m) of leaves (9.79 μmol·g^?1 DW h^?1 and 57.95 μm , respectively) were slightly higher than that of roots (6.09 μmol·g^?1DW h^?1 and 30.85 μm , respectively), whereas the affinity coefficients (α = V_max/K_m) for ammonium of leaves (0.17) and roots (0.20) were comparable, a characteristic that is unique among seagrass species. No substantial translocation (<2.5%) of ¹⁵N incorporated as ammonium was detected between plant parts, whereas the translocation of ¹⁵N incorporated as nitrate was higher (40–100%). We conclude that the N_i acquisition strategy of H. stipulacea, characterized by a similar uptake capacity and efficiency of leaves and roots, favors the geographical expansion potential of the species into areas with variable water‐sediment N levels throughout the Mediterranean.     

Sewage diversion effects on the water column of a subtropical estuary

A study of the phytoplankton community and water column chemistry in Kaneohe Bay, Oahu, Hawaii, before and after the diversion of secondary treated sewage from the bay has shown that changes in total nutrient concentrations in the water column cannot be accurately predicted without taking into account water column-benthos interactions. During the first year after sewage diversion, the decomposition of about 400 tonnes of benthic organisms, primarily filter feeders, resulted in water column dissolved organic nitrogen and phosphorus concentrations roughly an order of magnitude higher than those expected in the absence of such interactions. The biomass of phytoplankton appears to have been N limited both before and after sewage diversion, with internal nutrient recycling accounting for 70–99% of phtyoplankton nutrient uptake. Both the biomass and growth rate of the phytoplankton declined as a result of the sewage diversion, with post-diversion growth rates evidently well below nutrient-saturated values. Since the principal stresses on the bay's coral reef community as a result of the sewage discharges appear to have resulted from the elevated concentrations of plankton in the water, various measures of seston concentration appear to be the most ecologically significant indicators of nutrient enrichment in this system.