Photosynthesis and nitrogen fixation in a cyanobacterial bloom in the Baltic Sea

Daily integrals of photosynthesis by a cyanobacterial bloom in the Baltic Sea, during the summer of 1993, were calculated from the vertical distributions of light, temperature and the organisms in the water column and from photosynthesis/irradiance curves of picoplanktonic and diazotrophic cyanobacteria isolated from the community. The distribution of chlorophyll a in size-classes <20?µm and >20?µm was monitored over 9 days that included a deep mixing event followed by calm. Picocyanobacteria formed 70% of the cyanobacterial biomass and contributed 56% of the total primary production. Of the filamentous diazotrophs that formed the other 30%, Aphanizomenon contributed 28% and a Nodularia-containing fraction 16% of the primary production. For the whole population there was little change in standardized photosynthetic O₂ production, which remained at about 31?mmol?m^?2 before and after the mixing event. There were differences, however, between the classes of cyanobacteria: in picocyanobacteria primary production hardly changed, while in Aphanizomenon it increased by 2.6 and in Nodularia it fell below zero. Total phytoplankton photosynthesis was strongly dependent on total daily insolation with the compensation point at a photon insolation of 22.7?mol?m^?2?d^?1. Similar analyses of N₂ fixation showed much less dependence on depth distribution of light and biomass: Aphanizomenon fixed about twice as much N₂ as Nodularia their; their fixation exceeded their own N demand by about 12%. Together, these species contributed 49% of the total N demand of the phytoplankton population. Computer models based on the measured light attenuation and photosynthetic coefficients indicate that growth of the cyanobacterial population could occur only in the summer months when the critical depth of the cyanobacteria exceeds the depth of mixing.     

Nitrogen fixation by Baltic cyanobacteria is adapted to the prevailing photon flux density

N₂ fixation, measured as acetylene reduction, was studied in laboratory cultures and in natural assemblages (both as a mixed population and as individually picked colonies) of the heterocystous cyanobacteria Aphanizomenon sp. and Nodularia spp. from the Baltic Sea. During a diurnal cycle of alternating light and darkness, these organisms reduced acetylene predominantly during the period of illumination, although considerable activity was also observed during the dark period. In both laboratory cultures and natural populations N₂ fixation was saturated below a photon flux density of 600 μm⁻² s⁻¹. In cyanobacterial blooms in the Baltic Sea, nitrogenase activity was mostly confined to the surface layers. Samples collected from greater depths did not possess the same capacity for acetylene reduction as samples from the surface itself, even when incubated at the photon flux density prevailing in surface waters. This suggests that, with respect to N₂ fixation, Baltic cyanobacteria are adapted to the intensity of illumination that they are currently experiencing.     

Food web structure variability in the surface layer, at a fixed station influenced by the North Western Mediterranean Current

The structure of the planktonic community and the influence of mesozooplankton migration on the microbial food web were investigated during six diel studies from June 92 to June 93 in the surface waters of a station in the North-Western Mediterranean Sea. Each diel study consisted of sampling at 5 and 40 m every 3 h over 24 h. Most of the times diel cycles did not show any convincing diel patterns in any of the variables studied. Clear zooplankton migration was evident in only two diel studies.The ratio of heterotrophic/autotrophic biomasses varied from 0.68 to 3.0, with a strong dominance of the heterotrophic biomass under oligotrophic conditions. Differences in food web structure were probably related to the influences of coastal water and the North-Western Mediterranean Current. Thus we found that the planktonic food web variability relatable to hydrodynamic variability, to be greater than diel variability. However, very large differences in food web structure among dates were evident. For example proportion of Chl a found in the <10 µm fraction varied from 18 to 96%.     

Appearance and establishment of diazotrophic cyanobacteria in Lake Kinneret,Israel

1. We propose that the appearance and establishment of Nostocales (cyanobacteria) species of the genera Aphanizomenon and Cylindrospermopsis in the warm subtropical Lake Kinneret (Sea of Galilee, Israel) from 1994 was linked to changes in climate conditions and summer nitrogen (N) availability. 2. From 1994 to 2009, an increase in frequency of events of elevated water temperature (>29 °C) in summer, and to some extent a greater frequency of lower summer wind speed events, affected water turbulence and water column stratification, thus providing better physical conditions for the establishment of these populations. 3. In recent years, N‐depleted conditions in Lake Kinneret in early summer have promoted the development and domination of Nostocales that could gain an ecological advantage owing to their N₂‐fixing capability. 4. Nitrogen fixation rates coincided both with heterocyst abundance and with Nostocales biomass. The N supplied to the lake via nitrogen fixation ranged from negligible quantities when Nostocales represented only a minor component of the phytoplankton community to 123 tonnes when Cylindrospermopsis bloomed in 2005. This high N₂ fixation rate equals the average summer dissolved inorganic nitrogen load to the lake via the Jordan River.     

稳定性同位素技术在生态学上的应用

稳定性同位素技术早在20世纪70年代末期就被引入到生态学领域。最初是利用植物稳定性碳同位素的差异。开展了许多有关营养流动方面的研究;到90年代,稳定性碳和氮同位素被用来分析动物的食性、营养级位置关系以及食物链结构;本世纪初,由于技术的进步,稳定性同位素(特别是氢同位素)被用来开展动物迁徙习性方面的研究。到目前为止,国内有关这方面的研究还鲜有报道,而且对自然界存在的稳定性同位素的理解还存在一定偏差。本文主要介绍了稳定性同位素效应及其分馏原理、稳定性同位素在示踪动物食性信息、确定营养级位置关系、分析食物网结构以及研究动物迁徙生态学中的作用等方面的内容。     

Do top-down and bottom-up controls interact to exclude nitrogen-fixing cyanobacteria from the plankton of estuaries? An exploration with a simulation model

Explaining the nearly ubiquitous absence of nitrogen fixation by planktonic organisms in strongly nitrogen-limited estuaries presents a major challenge to aquatic ecologists. In freshwater lakes of moderate productivity, nitrogen limitation is seldom maintained for long since heterocystic, nitrogen-fixing cyanobacteria bloom, fix nitrogen, and alleviate the nitrogen limitation. In marked contrast to lakes, this behavior occurs in only a few estuaries worldwide. Primary production is limited by nitrogen in most temperate estuaries, yet no measurable planktonic nitrogen fixation occurs. In this paper, we present the hypothesis that the absence of planktonic nitrogen fixers from most estuaries is due to an interaction of bottom-up and top-down controls. The availability of Mo, a trace metal required for nitrogen fixation, is lower in estuaries than in freshwater lakes. This is not an absolute physiological constraint against the occurrence of nitrogen-fixing organisms, but the lower Mo availability may slow the growth rate of these organisms. The slower growth rate makes nitrogen-fixing cyanobacteria in estuaries more sensitive to mortality from grazing by zooplankton and benthic organisms.We use a simple, mechanistically based simulation model to explore this hypothesis. The model correctly predicts the timing of the formation of heterocystic, cyanobacterial blooms in freshwater lakes and the magnitude of the rate of nitrogen fixation. The model also correctly predicts that high zooplankton biomasses in freshwaters can partially suppress blooms of nitrogen-fixing cyanobacteria, even in strongly nitrogen-limited lakes. Further, the model indicates that a relatively small and environmentally realistic decrease in Mo availability, such as that which may occur in seawater compared to freshwaters due to sulfate inhibition of Mo assimilation, can suppress blooms of heterocystic cyanobacteria and prevent planktonic nitrogen fixation. For example, the model predicts that at a zooplankton biomass of 0.2 mg l^–1, cyanobacteria will bloom and fix nitrogen in lakes but not in estuaries of full-strength seawater salinity because of the lower Mo availability. Thus, the model provides strong support for our hypothesis that bottom-up and top-down controls may interact to cause the absence of planktonic nitrogen fixation in most estuaries. The model also provides a basis for further exploration of this hypothesis in individual estuarine systems and correctly predicts that planktonic nitrogen fixation can occur in low salinity estuaries, such as the Baltic Sea, where Mo availability is greater than in higher salinity estuaries.     

Effects of modified nutrient concentrations and ratios on the structure and function of the native phytoplankton community in the Neuse River Estuary, North Carolina, USA

A variety of analyses were used to assess the structure (community composition) and function (assimilation number, nitrogen fixation) of phytoplankton in the Neuse River Estuary (NRE), NC under ambient and modified nutrient concentrations. Dilution bioassays were employed to reduce the concentration of nitrogen (N) or both N and phosphorus (P) and thus compare varied DIN:DIP ratios. Experimental manipulations created conditions that may result from mandated N load reductions to the estuary. We hypothesized that unilateral reduction of N loading to the NRE would increase the activity, abundance and diversity of N₂ fixing cyanobacteria. Changes in phytoplankton primary productivity, N₂ fixation (nitrogenase activity), genetic potential for N₂ fixation (presence of nifH), phytoplankton taxonomic composition (diagnostic photopigment concentration) and abundances of N₂ fixing cyanobacteria (microscopy) were determined. Decreasing ambient DIN:DIP ratios in NRE samples resulted in increased rates of N₂ fixation when seed populations were present and environmental conditions were amenable. Decreasing the DIN:DIP ratio did not lead to an increase in the abundance or diversity of N₂ fixing cyanobacteria. Because N₂ fixing cyanobacteria were only actively fixing nitrogen during periods of low riverine N discharge (summer and early autumn), lowering nutrient ratios may not have a major impact on the NRE. However, the maximum potential amount of N from N₂ fixation was calculated using rates from this study and was found to be approximately 3% of total riverine loading of N to the NRE. Because N₂ fixation occurs farther downstream and later in the year than riverine N loading to the NRE, there is potential for N₂ fixation to modify N dynamics. Analyses of the phytoplankton community as a whole in these relatively short term experiments indicated that reduced DIN:DIP may not have a major impact on their structure and function.     

应用碳,氮稳定同位素研究稻田多个物种共存的食物网结构和营养级关系

稳定碳、氮同位素比值分析技术是研究生态系统中物质循环与能量流动的有效技术。δ13C值常用来分析消费者食物来源,δ15N值常用来确定生物在食物网中的营养位置。应用稳定同位素技术分析了稻-鱼(R-F)和稻-鱼-鸭(R-F-D)两种稻作方式下,稻田多个物种共存的食物网结构和营养级关系。结果表明,R-F中SOM的δ13C值为(-27.7±0.3)‰,与R-F-D(-27.4±0.4)‰,相差不大;R-F中POM的1δ3C值为(-27.4±0.8)‰,低于稻鱼鸭共生田(-26.7±0.5)‰;δ15N值计算发现R-F内浮游动物的营养级位置在2.24±0.16,鱼的营养级位置在3.07±0.26,均高于其在R-F-D内的营养级。在R-F-D内,由于鸭的引入,和R-F相比,鱼的营养级降低为2.63±0.13。     

Nitrogen Fixation During Blooms of Nodularia in Coastal Waters and Backwaters of the Arkona Sea (Baltic Sea) in 1974

Nitrogen fixation was investigated by means of the acetylene reduction method during the development of a water bloom of Nodularia in coastal waters of the Baltic Sea west of the island of Hiddensee and in backwaters showing different degrees of eutrophication. Depending on plankton density, the values found varied greatly. The maximum of nitrogen fixation values found in extremely dense water blooms under special conditions (Baltic Sea, 2250 μg N₂/l · h; Kleiner Jasmunder Bodden, 374 μg N₂/l · h) are up to 10³ times higher than from other parts of the Baltic Sea or from inland waters. The average nitrogenase activity determined for coastal water populations of the Baltic Sea is 2.15 pg N₂/heterocyst · h and that of the inmost backwaters 0.77 pg N₂/heterocyst · h. The relationship between N₂-fixation and nutrient content in water is discussed.     

Amino acid availability determines the ratio of microcystin variants in the cyanobacterium Planktothrix agardhii

Cyanobacteria are capable of producing multiple microcystin variants simultaneously. The mechanisms that determine the composition of microcystin variants in cyanobacteria are still debated. [Asp³]microcystin-RR contains arginine at the position where the more toxic [Asp³]microcystin-LR incorporates leucine. We cultured the filamentous cyanobacterium Planktothrix agardhii strain 126/3 with and without external addition of leucine and arginine. Addition of leucine to the growth medium resulted in a strong increase of the [Asp³]microcystin LR/RR ratio, while addition of arginine resulted in a decrease. This demonstrates that amino acid availability plays a role in the synthesis of different microcystin variants. Environmental changes affecting cell metabolism may cause differences in the intracellular availability of leucine and arginine, which can thus affect the production of microcystin variants. Because leucine contains one nitrogen atom while arginine contains four nitrogen atoms, we hypothesized that low nitrogen availability might shift the amino acid composition in favor of leucine, which might explain seasonal increases in the [Asp³]microcystin LR/RR ratio in natural populations. However, when a continuous culture of P. agardhii was shifted from nitrogen-saturated to a nitrogen-limited mineral medium, leucine and arginine concentrations decreased, but the leucine/arginine ratio did not change. Accordingly, while the total microcystin concentration of the cells decreased, we did not observe changes in the [Asp³]microcystin LR/RR ratio in response to nitrogen limitation.     

Spatial heterogeneity of trophic pathways in the invertebrate community of a temperate bog

Summary 1. To examine spatial heterogeneity of trophic pathways on a small scale (<5 m diameter), we conducted dual stable isotope (δ¹³C and δ¹⁵N) analyses of invertebrate communities and their potential food sources in three patchy habitats [sphagnum lawn (SL), vascular‐plant carpet (VC) and sphagnum carpet] within a temperate bog (Mizorogaike Pond, Kyoto, Japan). 2. In total, 19 invertebrate taxa were collected from the three habitats, most of which were stenotopic, i.e. collected from a single habitat. Amongst the habitats, significant variation was observed in the isotopic signatures of dominant plant tissues and their detrital matter [benthic particulate organic matter (BPOM)], both of which were potential organic food sources for invertebrates. Site‐specific isotopic variation amongst detritivores was found in δ¹³C but not in δ¹⁵N, reflecting site‐specificity in the isotopic signatures of basal foods. The eurytopic hydrophilid beetle Helochares striatus was found in all habitats, but showed clear site variation in its isotopic signatures, suggesting that it strongly relies on foods within its own habitat. 3. The most promising potential foods for detritivores were the dead leaf stalks of a dominant plant in the VC and BPOM in the SL and carpet. An isotopic mixing model (IsoSource version 1.3.1) estimated that aquatic predators rely on unknown trophic sources with higher δ¹³C than detritus, whereas terrestrial predators forage on allochthonous as well as autochthonous prey, suggesting that the latter predators might play key roles in coupling between habitats. 4. Our stable isotope approach revealed that immobile detritivores are confined to their small patchy habitats but that heterogeneous trophic pathways can be coupled by mobile predators, stressing the importance of habitat heterogeneity and predator coupling in characterising food webs in bog ecosystems.