Do cyanobacteria dominate in eutrophic lakes because they fix atmospheric nitrogen?

1. The sources of nitrogen for phytoplankton were determined for a bloom‐prone lake as a means of assessing the hypothesis that cyanobacteria dominate in eutrophic lakes because of their ability to fix nitrogen when the nitrogen : phosphorous (N : P) supply ratio is low and nitrogen a limiting resource. 2. Nitrogen fixation rates, estimated through acetylene reduction with ¹⁵N calibration, were compared with ¹⁵N‐tracer estimates of ammonium and nitrate uptake monthly during the ice‐free season of 1999. In addition, the natural N stable isotope composition of phytoplankton, nitrate and ammonium were measured biweekly and the contribution of N₂ to the phytoplankton signature estimated with a mixing model. 3. Although cyanobacteria made up 81–98% of phytoplankton biomass during summer and autumn, both assays suggested minimal N acquisition through fixation (<9% for the in‐situ incubations; <2% for stable isotope analysis). Phytoplankton acquired N primarily as ammonium (82–98%), and secondarily as nitrate (15–18% in spring and autumn, but <5% in summer). Heterocyst densities of <3 per 100 fixer cells confirmed low reliance on fixation. 4. The lake showed symptoms of both light and nitrogen limitation. Cyanobacteria may have dominated by monopolizing benthic sources of ammonium, or by forming surface scums that shaded other algae.     

Carbon, nitrogen, and phosphorus dynamics during leaf decay in nutrient-enriched stream microecosystems

We investigated the effects of long-term enrichment with nitrate, phosphate, and nitrate+phosphate on the first 5 weeks of leaf detritus processing in laboratory stream microecosystems. Enrichment with nitrate+phosphate accelerated leaf weight loss and increased rates of respiration associated with the leaves. However, whole-system respiration was little changed from that observed in the control stream since respiration in the water was greatly reduced. Enrichment with phosphate alone had little effect except to lower respiration associated with leaf discs. Enrichment with nitrate alone also decreased leaf-disc respiration but resulted in a greatly increased rate of respiration in the water. Net leaching and fragmentation of carbon from the leaves was also increased by nitrate enrichment. Nitrogen and phosphorus levels in leaf material were little affected by enrichment with nitrate or phosphorus alone. Leaves in those streams and in the control stream released nitrogen and phosphorus to the water. In contrast, percent nitrogen and phosphorus increased greatly in the leaves in the stream enriched with both nitrate and phosphate. The leaves in this system immobilized both nitrogen and phosphorus from the water. We also studied the importance of nitrogen fixation as a vector for nitrogen incorporation associated with leaf decomposition in streams. Somewhat surprisingly, fixation by microbes associated with the leaves and by microbes suspended in the water occurred under all three experimental enrichment treatments as well as in the control, casting doubt on the effectiveness of nitrate in inhibiting nitrogenase synthesis in nature. However, N₂-fixation is only a minor source of nitrogen for leaves decaying under the conditions studied.     

Population responses of Lake Michigan phytoplankton to nitrogen and phosphorus enrichment

A natural phytoplankton assemblage from Grand Traverse Bay, Lake Michigan, was treated with factorial enrichments of nitrate and phosphorus, with maintained nutrient concentrations ranging from 5 to 60 μg total soluble phosphorus liter⁻¹ and 0.225 to 1.12 mg nitrate-nitrogen liter⁻¹.One container was spiked with added vitamins, a chelator, and trace metals. The assemblage response was monitored at the species level. Significant differences in growth rates as a function of nutrient enrichment were detected at both the division and the species levels. Growth rates associated with the various levels of enrichment are reported for several diatom taxa. Many of the diatom taxa exhibited highly significant (P < 0.01) increases in growth rate after phosphorus enrichment, with the largest effects occurring between 5 and 15 μg total soluble phosphorus liter⁻¹. Significant (P < 0.05) N effects were also observed, and the nature of these effects was found to be taxon-specific. Taxa also showed significant changes in percent composition, due both to time and to nutrient enrichment, indicating a substantial heterogeneity in response at the species level. Experimentally induced population changes were qualitatively similar to those observed in regions of the Great Lakes which have undergone anthropogenic eutrophication. Contribution No. 222 of the Great Lakes Research Division, University of Michigan. Work was supported by funds from the Michigan Sea Grant Program and the Environmental Protection Agency. Contribution No. 222 of the Great Lakes Research Division, University of Michigan. Work was supported by funds from the Michigan Sea Grant Program and the Environmental Protection Agency.     

The response of planktonic and microbenthic algal assemblages to nutrient enrichment in shallow coastal waters,southwest Sweden

Field and laboratory nutrient (nitrogen and phosphorus) enrichment experiments were performed using natural phytoplankton and microphytobenthic assemblages from the brackish water Öresund, S.W. Sweden. The response of algae from a low-nutrient area (Falsterbo Canal) was compared to that of algae from a polluted, nutrient-rich area (Lomma Bay).The biomass (measured as chlorophyll a) of both phytoplankton and microphytobenthos from the Falsterbo Canal increased after the addition of nitrogen. Phytoplankton growth was stimulated by the addition of phosphorus to the nitrogen-rich water of the polluted Lomma Bay. Sediment chlorophyll a showed no significant increase after the addition of nutrients in the Lomma Bay. In containers without sediment, phytoplankton uptake was calculated to account for ≈ 90% of the disappearance of inorganic fixed nitrogen from the water. In the sediment containers the microphytobenthos was estimated to account for ≈20% of the nitrogen uptake. The rest was presumably lost mainly through denitrification.When containers with microphytobenthos from Lomma Bay were kept in the dark, phosphorus was released at a rate of up to ≈ 180 μM · m^?2 · day^?1. We suggest that by producing oxygen microbenthic algae keep the sediment surface oxygenated thereby decreasing phosphorus transport from the sediment to the overlying water.     

Monitoring plant physiological characteristics to evaluate mine site revegetation: A case study from the wet-dry tropics of northern Australia

Biologically driven markers or monitors were used to evaluate plant and ecosystem health of uranium-mining affected sites. Plant water, nitrogen (N) and phosphorus (P) status were used to measure physiological characteristics of tree and shrub species at sites perturbed by mining activities (waste rock dumps: WRD 1, WRD 2; mine wastewater irrigated woodland) and of species at undisturbed woodland (tropical savanna). Plant water status was evaluated by measuring leaf relative water content (RWC) and carbon isotope discrimination (δ¹³C). Leaf RWC varied significantly (P<0.0001) between wet and dry season in species at the woodland sites with higher RWC in the wet season compared to the dry season. No seasonal differences were observed in RWC in species at the WRDs. Leaf δ¹³C was similar in species at woodland sites and WRD 2 (−28.8 to −28.1‰) but was significantly (P<0.05) lower in species at WRD 1 (−27.6‰). This suggests that species at WRD 1 had a lower water availability and/or lower water use compared to species at all other sites. WRD substrate had an up to 4-orders of magnitude greater availability of inorganic phosphate (Pi) compared to woodland soil as determined using in situ ion exchange resin. Pi concentrations in xylem sap of species at WRDs were 2- to 3-fold higher compared to species at woodland sites. Plant nitrate reductase (NR) activity was low in most species at woodland and WRD 1. In contrast, Eucalyptus and Acacia species had high NR activities of up to 300–700 pkat g^-1 fw at WRD 2 indicating that these species had greater nitrate use than species at all other sites. Nitrate availability in the top five cm of the profile, as determined using in situ ion exchange resins, increased at all sites in the wet season, but no significant differences were observed between sites using this method. However, traditional soil analysis revealed that WRD substrate had a 2-times higher nitrate content (0 to 1000 mm depth) compared to woodland soil. Thus, it is likely that plants at WRD2 accessed nitrate from deeper parts of the profile. Proline, an indicator of plant stress, was found in appreciable quantities in leaves of herbaceous species but not in woody species. Soil and leaf δ¹⁵N were measured to investigate N-cycling and the contribution of diazotrophic N₂ fixation to plant N nutrition. Soil δ¹⁵N values were highest and most variable at WRD 2 (6.2‰) compared to all other sites (irrigated woodland 3.1‰, undisturbed woodland 2.5‰, WRD 1 0.9‰). This may indicate that N-turnover and nitrification was greatest at WRD 2 leading to greater ¹⁵N enrichment of soil N. At all sites, Acacia species were nodulated and putatively fixing N₂. With the exception of WRD 2 where leaf δ¹⁵N of Acacia species averaged 0.9‰, Acacia species had ¹⁵N depleted values characteristic of species that receive N derived from N₂ fixation (−0.8 to −0.6‰). Eucalyptus species at the woodland also had ¹⁵N depleted values (average −0.4‰) but ¹⁵N enriched values (0.3 to 1.8‰) at the three mining affected sites. The results show that for the plants studied foliar δ¹⁵N could not be used as an unequivocal measure of plant N sources. The results suggest that biomonitoring of plant and ecosystem health has potential in evaluating performance of mine site revegetation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Nutrient limitation of phytoplankton growth in Waquoit Bay, Massachusetts, USA: a nutrient enrichment study

We conducted nutrient enrichment experiments and field sampling to address three questions: (1) is there nutrient limitation of phytoplankton accumulation within an estuary whose waters are exposed to relatively high nitrogen loading rates, (2) where in the salinity gradient from fresh to seawater (0 to 32‰) is there a shift from phosphorus to nitrogen limitation of phytoplankton accumulation, and (3) is there a seasonal shift in limiting function of phosphorus and nitrogen anywhere in the estuarine gradient. Nitrogen and phosphorus enrichment experiments in the Childs River, an estuary of Waquoit Bay, Massachusetts, USA, showed that the accumulation of phytoplankton biomass in brackish and saline water was limited by supply of nitrate during warm months. The effects of enrichment were less evident in fresh water, with short-lived responses to phosphate enrichment. There was no specific point along the salinity gradient where there was a shift from phosphorus- to nitrogen-limited phytoplankton accumulation; rather, the relative importance of nitrogen and phosphorus changed along the salinity gradient in the estuary and with season of the year. There was no response to nutrient additions during the colder months, suggesting that some seasonally-varying factor, such as light, temperature or a physiological mechanism, restricted phytoplankton accumulation during months other than May-Aug. There was only slight evidence of a seasonal shift between nitrogen- and phosphorus-limitation of chlorophyll accumulation. Phytoplankton populations in nutrient-rich estuaries with short flushing times grow fast, but at the same time the cells may be advected out of the estuaries while still rapidly dividing, thereby providing an important subsidy to production in nearby deeper waters. This revised version was published online in August 2006 with corrections to the Cover Date.     

Impacts of invading N2-fixing Acacia species on patterns of nutrient cycling in two Cape ecosystems: evidence from soil incubation studies and 15N natural abundance values

This study examines the impacts of woody, N₂-fixing invasive Acacia spp. on the patterns of nutrient cycling in two invaded ecosystems of differing nutrient status in the Cape floristic region. Patterns of soil nutrient mineralization were measured by a field incubation method while the significance of the fixation process in altering nutrient cycling was assessed by the ¹⁵N natural abundance technique. The results confirm earlier reports that invasion by woody shrubs results in organic matter and nutrient enrichment of surface soils of both ecosystems. However, patterns of nutrient availability (phosphorus and nitrogen) were not necessarily enhanced. In the more fertile strandveld both phosphorus and nitrogen (significant at P<0.10) showed trends towards enhanced annual mineralization rates upon invasion, while in the low nutrient fynbos system only phosphorus followed this trend. It is unclear whether this differential response is a consequence of plant- or soil-derived feedbacks on the decomposition processes in each system. The ¹⁵N values of the soils from the invaded sites of both ecosystems indicated a strong influence of the alien species on the soil nitrogen component. However, as with other studies of natural ecosystems, the contribution of nitrogen from fixation could not be readily quantified with the ¹⁵N natural abundance method because of problems in selecting suitable non-N₂-fixing reference plants. A technique of disrupting nodule structure and function, by fumigation with O₂, to obtain the ¹⁵N value of a non-N₂-fixing speciment of the study species was tried and found to overcome some of the problems associated with the lack of suitable reference plants. With this technique it was possible to detect the almost total dependence of A. saligna on N₂-fixation in the fynbos soils with their low nitrogen mineralization rates. In the strandveld ecosystem with much higher soil nitrogen release rates A. cyclops was only partly dependent on fixation (about half) for its nitrogen. The nutrient enrichment of both ecosystems and trends towards enhanced rates of nutrient mineralization could have profound implications on the long-term success of alien invader clearing operations and the restoration of the indigenous flora at these sites.     

Nutrient limitation of phytoplankton and periphyton growth in upland lakes

SUMMARY 1. Thirty small upland lakes in Cumbria, Wales, Scotland and Northern Ireland were visited three times between April and August 2000. On each occasion water chemistry was measured and phytoplankton bioassays were performed in the laboratory to assess growth‐rate and yield limitation by phosphorus and nitrogen. In addition, yield limitation of periphyton growth was investigated twice, in situ, using nutrient‐diffusing substrata. 2. Over the whole season the percentage frequency of P, N and co‐limitation was 24, 13 and 63%, respectively, for phytoplankton rate limitation and 20, 22 and 58%, respectively, for phytoplankton yield limitation. 3. A clear response of periphyton yield to nutrient additions was found in 75% of all cases and of these, co‐limitation was most common (54%). Average percentage frequency for P and N limitation was 26 and 20%, respectively. 4. Phytoplankton and periphyton showed seasonal changes in nutrient limitation within sites. In particular, co‐limitation became progressively more common as the season progressed. 5. The response of phytoplankton growth rate to ammonium and nitrate addition was identical, but ammonium was a slightly better source of nitrogen than nitrate for phytoplankton yield on 7% and for periphyton yield on 10% of the occasions. However, the magnitude of the effect was small. 6. The concentration of dissolved inorganic nitrogen (DIN) and the molar ratio of DIN to total dissolved phosphorus (TDP), appeared to be the main environmental factors controlling the extent of nitrogen or phosphorus limitation at a given site. Nitrogen limitation was more likely than phosphorus limitation where the DIN was <6.5 mmol m^?3 and the ratio of DIN : TDP was <53. Co‐limitation was the most likely outcome at a DIN concentration <13 mmol m^?3 and at a DIN : TDP molar ratio <250. Above these values phosphorus limitation was most likely. 7. The relatively high frequency of nitrogen limitation and co‐limitation at higher N : P ratios than previously reported, may result from the inability of nitrogen‐fixing cyanobacteria to thrive in these upland lakes where pH and the concentration of phosphorus tended to be low and where flushing rates tended to be high.     

Nitrogen enrichment during decomposition of mangrove leaf litter in an east African coastal lagoon (Kenya): Relative importance of biological nitrogen fixation

In situ decomposition of senescent leaves of twoabundant mangrove species (Rhizophora mucronataLamarck and Ceriops tagal (Perr) C.B. Rob),enrichment of nitrogen and activity of dinitrogenfixing bacteria during decomposition were investigatedduring both rainy and dry seasons in a tropicalcoastal lagoon (Gazi, Kenya). Rates of leafdecomposition were higher for R. mucronata thanfor C. tagal and were highest, for both species,during rainy season. Rates of decomposition, expressedas percentage dry mass loss, over a decompositionperiod of 50 days was: C. tagal (rainy season),69%; C. tagal (dry season), 27%; R.mucronata (rainy season), 98%; and R.mucronata (dry season), 48%. High rainfall anddiurnal tidal inundation appear to enhance the leafdecomposition process. Maximum rates of nitrogenfixation were 380 nmol N₂ h^-1 g^-1 dw forC. tagal (rainy season), 78 nmolN₂ h^-1 g^-1 dw for C. tagal (dryseason), 390 nmol N₂ h^-1 g^-1 dw for R. mucronata (rainy season) and 189 nmolN₂ h^-1 g^-1 dw for R. mucronata (dry season). Although N₂ fixation rates werehighest during rainy season, total nitrogenimmobilised in the leaves was highest during the dryseason. Biological nitrogen fixation can account forbetween 13 to 21% of the maximum nitrogen immobilisedin the decaying mangrove leaves. Nitrogen fixation, asa source of allochthonous nitrogen, sustains anitrogen input to the mangrove ecosystem, which addssignificantly to the nitrogen input through leaflitterfall.     

Nitrogen dynamics and microbial food web structure during a summer cyanobacterial bloom in a subtropical,shallow, well-mixed,eutrophic lake (Lake Taihu,China)

Nitrogen dynamics and microbial food web structure were characterized in subtropical, eutrophic, large (2,338 km²), shallow (1.9 m mean depth), and polymictic Lake Taihu (China) in Sept–Oct 2002 during a cyanobacterial bloom. Population growth and industrialization are factors in trophic status deterioration in Lake Taihu. Sites for investigation were selected along a transect from the Liangxihe River discharge into Meiliang Bay to the main lake. Water column nitrogen and microbial food web measurements were combined with sediment–water interface incubations to characterize and identify important processes related to system nitrogen dynamics. Results indicate a gradient from strong phosphorus limitation at the river discharge to nitrogen limitation or co-limitation in the main lake. Denitrification in Meiliang Bay may drive main lake nitrogen limitation by removing excess nitrogen before physical transport to the main lake. Five times higher nutrient mineralization rates in the water column versus sediments indicate that sediment nutrient transformations were not as important as water column processes for fueling primary production. However, sediments provide a site for denitrification, which, along with nitrogen fixation and other processes, can determine available nutrient ratios. Dissimilatory nitrate reduction to ammonium (DNRA) was important, relative to denitrification, only at the river discharge site, and nitrogen fixation was observed only in the main lake. Reflecting nitrogen cycling patterns, microbial food web structure shifted from autotrophic (phytoplankton dominated) at the river discharge to heterotrophic (bacteria dominated) in and near the main lake.     

Comparison of acetylene-reduction and nitrogen-15 techniques for the determination of nitrogen fixation by field bean (Vicia faba) nodules

Field bean (Vicia faba L.) cv. Maris Bead seeds were inoculated with Rhizobium Catalogue No. 1001, supplied by Rothamsted Experimental Station, and grown in sand culture supplied with a complete nutrient solution which included nitrate at either 1.5 or 6.0 mM. Nodules were detached from the roots at intervals during plant development and their rates of nitrogen fixation estimated by both acetylene reduction and ¹⁵N gas technique. There was a constant relationship, independent of nitrate supply, between the results obtained by these two methods at all samplings. The amounts of acetylene reduced divided by a factor of 5.75 gave the amount of true nitrogen fixation; this factor is about twice the theoretical value. It is suggested that this discrepancy arose because, with acetylene, all the electrons available to the nitrogenase were used to form ethylene, whereas during normal fixation only about half the electron supply was used to fix nitrogen, the remainder having been consumed in the production of hydrogen gas.     

The role of heterocytes in the physiology and ecology of bloom-forming harmful cyanobacteria

Dolichospermum flos-aquae and Cylindrospermopsis raciborskii are two cyanobacteria species which cause harmful blooms around the world. Both these species share the capacity to fix atmospheric nitrogen in heterocytes (cell where fixation occurs). While Dolichospermum can express heterocytes at rather regular intervals across the filament, Cylindrospermopsis can only express heterocytes at the end of the filament. The aim of this study was to experimentally assess the role of heterocyte position in the eco-physiological responses of these bloom forming cyanobacteria. Replicated monocultures of each species were grown at different eutrophication scenarios (limiting and sufficient nitrogen and phosphorus concentrations, in factorial design). Dolichospermum reached high biomass regardless of the nitrogen (and phosphorus) provided, suggesting that this species could bloom in situations with and without nitrogen limitation. In contrast, Cylindrospermopsis reached high biomass only when nitrogen supply was high; its biomass was 15-20 times lower when relying on nitrogen fixation. Hence, despite its ability to fix nitrogen, blooms of Cylindrospermopsis would be expected only under high total nitrogen availability. In Dolichospermum heterocytes occurred only in the scenarios without supplied nitrogen while in Cylindrospermopsis heterocytes occurred regardless of nitrogen availability. Yet, in both species nitrogen fixation occurred (heterocytes were functional) only when nitrogen was limiting, and nitrogen fixation increased significantly at higher phosphorus concentration. Finally, in the absence of supplied nitrogen, filament length in Dolichospermum was the longest, while filaments in Cylindrospermopsis were the shortest (up to 13 times shorter than at nitrogen sufficiency). Therefore, heterocyte expression in Dolichospermum, and filament length in Cylindrospermopsis seem good proxies of nitrogen fixation. The eco-physiological responses recorded here help understand the distribution of these species along nutrient gradients in nature.     

Nutrient Biogeochemistry During the Early Stages of Delta Development in the Mississippi River Deltaic Plain

Nutrient biogeochemistry associated with the early stages of soil development in deltaic floodplains has not been well defined. Such a model should follow classic patterns of soil nutrient pools described for alluvial ecosystems that are dominated by mineral matter high in phosphorus and low in carbon and nitrogen. A contrast with classic models of soil development is the anthropogenically enriched high nitrate conditions due to agricultural fertilization in upstream watersheds. Here we determine if short-term patterns of soil chemistry and dissolved inorganic nutrient fluxes along the emerging Wax Lake delta (WLD) chronosequence are consistent with conceptual models of long-term nutrient availability described for other ecosystems. We add a low nitrate treatment more typical of historic delta development to evaluate the role of nitrate enrichment in determining the net dinitrogen (N₂) flux. Throughout the 35-year chronosequence, soil nitrogen and organic matter content significantly increased by an order of magnitude, whereas phosphorus exhibited a less pronounced increase. Under ambient nitrate concentrations (>60 μM), mean net N₂ fluxes (157.5 μmol N m^?2 h^?1) indicated greater rates of gross denitrification than gross nitrogen fixation; however, under low nitrate concentrations (<2 μM), soils switched from net denitrification to net nitrogen fixation (?74.5 μmol N m^?2 h^?1). As soils in the WLD aged, the subsequent increase in organic matter stimulated net N₂, oxygen, nitrate, and nitrite fluxes producing greater fluxes in more mature soils. In conclusion, soil nitrogen and carbon accumulation along an emerging delta chronosequence largely coincide with classic patterns of soil development described for alluvial floodplains, and substrate age together with ambient nitrogen availability can be used to predict net N₂ fluxes during early delta evolution.     

Hydrological and nutrient budgets of freshwater and estuarine wetlands of Taylor Slough in Southern Everglades, Florida (U.S.A.)

Hydrological restoration of the Southern Everglades will result in increased freshwater flow to the freshwater and estuarine wetlands bordering Florida Bay. We evaluated the contribution of surface freshwater runoff versus atmospheric deposition and ground water on the water and nutrient budgets of these wetlands. These estimates were used to assess the importance of hydrologic inputs and losses relative to sediment burial, denitrification, and nitrogen fixation. We calculated seasonal inputs and outputs of water, total phosphorus (TP) and total nitrogen (TN) from surface water, precipitation, and evapotranspiration in the Taylor Slough/C-111 basin wetlands for 1.5 years. Atmospheric deposition was the dominant source of water and TP for these oligotrophic, phosphorus-limited wetlands. Surface water was the major TN source of during the wet season, but on an annual basis was equal to the atmospheric TN deposition. We calculated a net annual import of 31.4 mg m^–2 yr^–1 P and 694 mg m^–2 yr^–1N into the wetland from hydrologic sources. Hydrologic import of P was within range of estimates of sediment P burial (33–70 mg m^–2 yr^–1 P), while sediment burial of N (1890–4027 mg m^–2 yr^–1 N) greatly exceeded estimated hydrologic N import. High nitrogen fixation rates or an underestimation of groundwater N flux may explain the discrepancy between estimates of hydrologic N import and sediment N burial rates.     

Whole-stream nitrate addition affects litter decomposition and associated fungi but not invertebrates

We assessed the effect of whole-stream nitrate enrichment on decomposition of three substrates differing in nutrient quality (alder and oak leaves and balsa veneers) and associated fungi and invertebrates. During the 3-month nitrate enrichment of a headwater stream in central Portugal, litter was incubated in the reference site (mean NO₃-N 82 μg l⁻¹) and four enriched sites along the nitrate gradient (214–983 μg NO₃-N l⁻¹). A similar decomposition experiment was also carried out in the same sites at ambient nutrient conditions the following year (33–104 μg NO₃-N l⁻¹). Decomposition rates and sporulation of aquatic hyphomycetes associated with litter were determined in both experiments, whereas N and P content of litter, associated fungal biomass and invertebrates were followed only during the nitrate addition experiment. Nitrate enrichment stimulated decomposition of oak leaves and balsa veneers, fungal biomass accrual on alder leaves and balsa veneers and sporulation of aquatic hyphomycetes on all substrates. Nitrate concentration in stream water showed a strong asymptotic relationship (Michaelis–Menten-type saturation model) with temperature-adjusted decomposition rates and percentage initial litter mass converted into aquatic hyphomycete conidia for all substrates. Fungal communities did not differ significantly among sites but some species showed substrate preferences. Nevertheless, certain species were sensitive to nitrogen concentration in water by increasing or decreasing their sporulation rate accordingly. N and P content of litter and abundances or richness of litter-associated invertebrates were not affected by nitrate addition. It appears that microbial nitrogen demands can be met at relatively low levels of dissolved nitrate, suggesting that even minor increases in nitrogen in streams due to, e.g., anthropogenic eutrophication may lead to significant shifts in microbial dynamics and ecosystem functioning. Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

太湖西北湖区2003-2012年间氮磷浓度及浮游植物主要类群变化趋势分析

基于2003-2012年太湖竺山湖和西部沿岸区水体理化指标与浮游植物丰度的逐月监测数据,分析了两个湖区氮磷营养盐状态和浮游植物丰度以及浮游植物主要类群的年际变化趋势及季节变化特征,探讨了浮游植物群落变化与水温及营养盐指标间的关系。结果表明：10年间两个湖区氮磷营养盐浓度总体呈下降趋势,以竺山湖TN、NH₃-N浓度和西部沿岸区NO₃-N浓度下降最为显著;浮游植物丰度总体呈上升趋势,蓝藻在群落结构中日益占据绝对优势;季节变化上,氮营养盐浓度表现为春冬季节高于夏秋季节,TP浓度和浮游植物丰度呈相反的变化趋势。Pearson相关分析显示,水温、NH₃-N浓度和TN/TP是影响蓝藻丰度及其在浮游植物群落中所占比例的主要因素。在温度和营养盐结构的共同作用下,10年间两个湖区蓝藻水华暴发时间逐渐提前,而消退时间逐渐滞后,水华持续时间逐年上升。在全球变暖背景下,太湖水华治理需执行更加严格的氮磷限制阈值,且在重污染的西北湖区控磷依然是关键。     

Experiments on Nitrogen and Phosphorus Release by Chironomus ex gr. plumosus from the Sediments of Lake Balaton,Hungary

Sediment-water systems of Lake Balaton mud and tap water were constructed in plexiglass tubes and boxes for evaluation of changes in the nitrogen and phosphorus contents of overlying water as influenced by chironomid density and light conditions. Nitrogen flux from sediment was greatly influenced by denitrification process. The amount of TN released by chironomids amounted to about half of the yearly nitrogen fixation rate in Lake Balaton. Omnivorous chironomids increased the phosphorus content of the overlying water. The rate of release was density dependent. TP release rates ranged between 1–17 mg m⁻² day ⁻¹ over a range of densities 500–20,000 larvae m⁻². It became evident that aerobic sediment cores can be an important nutrient source in lakes where chironomids inhabit them at densities above 1,000 larvae m⁻².     

15N natural abundance of plant and soil components of a Banksia woodland ecosystem in relation to nitrate utilization, life form, mycorrhizal status and N2-fixing abilities of component species

Studies of the variation in δ¹⁵N values for plants from a fire-prone Banksia woodland in South West Australia showed that pioneer herbaceous, non-mycorrhizal species which were active in nitrate reduction and storage, had the highest values (1.81%c). A detailed study of one such species Ptilotus polystachus demonstrated a close correspondence between the δ¹⁵N values of soil nitrate, xylem nitrate and leaf total nitrogen, suggesting an exclusive reliance on nitrate ions as nitrogen source. These pioneer species also showed a preponderance of the chloroplastic isoform of glutamine synthetase while woody species generally had higher activity associated with the cytosolic isoform. The group comprising monocotyledonous hemicryptophytes and geophytes contained species with slightly positive δ¹⁵N values and moderately active in nitrate reduction and storage. Nitrogen-fixing species had the lowest δ¹⁵N values (–0.36‰), irrespective of their apparent utilisation of nitrate. However, woody resprouter species which had low levels of nitrate reduction and storage had δ¹⁵N values which fell within the range of values obtained for the miscellaneous assemblage of N₂-fixing species. Consequently, ¹⁵N abundance values failed to distinguish N₂ fixing from non-fixing woody species, and therefore, could not be used in the ecosystem to determine the dependence of putative nitrogen fixing species on N₂ fixation. The study demonstrated complex patterns of nitrogen utilization in the ecosystem in which exploitation of different nitrogen resources related to plant life form and the physiological attributes of nitrogen assimilation by component species.     

Evidence that Northern Pioneering Pines with Tuberculate Mycorrhizae are Unaffected by Varying Soil Nitrogen Levels

Tuberculate mycorrhizae on Pinus contorta (lodgepole pine) have previously been shown to reduce acetylene, but an outstanding question has been to what degree these structures could meet the nitrogen requirements of the tree. We compared the growth, tissue nitrogen contents, and stable nitrogen isotope ratios of P. contorta growing in gravel pits to the same species growing on adjacent intact soil. Trees growing in severely nitrogen deficient gravel pits had virtually identical growth rates and tissue nitrogen contents to those growing on intact soil that had nitrogen levels typical for the area. δ¹⁵N values for trees in the gravel pits were substantially lower than δ¹⁵N values for trees on intact soil, and isotope ratios in vegetation were lower than the isotope ratios of the soil. The form of soil nitrogen in the gravel pits was almost exclusively nitrate, while ammonium predominated in the intact soil. Discrimination against ¹⁵N during plant uptake of soil nitrate in the highly N-deficient soil should be weak or nonexistent. Therefore, the low δ¹⁵N in the gravel pit trees suggests that trees growing in gravel pits were using another nitrogen source in addition to the soil. Precipitation-borne nitrogen in the study area is extremely low. In conjunction with our other work, these findings strongly suggests that P. contorta and its microbial symbionts or associates fix nitrogen in sufficient amounts to sustain vigorous tree growth on the most nitrogen-deficient soils.     

氮输入对植物光合固碳的影响研究进展

植物光合固碳(C)是生物固C的重要途径和生态系统C循环中的重要环节。在全球环境变化背景下,研究氮(N)输入对植物光合固C的影响,对于更好的认识C、N循环过程及生态系统对全球变化的响应过程等具有重要意义。N输入是否能够增加植物固C取决于生态系统类型以及生态系统的N饱和度;草原和湿地生态系统N输入的临界负荷值较高,干旱、半干旱荒漠地区较低;N输入可能改变植物光合固C在各器官的分配,主要由植物生理、自身生长节律和环境养分等决定。由于物种和生态系统类型的差异,N输入对植物固C的影响仍具有很大的不确定性,目前缺乏准确、定量表达N输入对生态系统光合和C同化物分配影响的数学表达方法和过程算法。未来应着重加强N输入下C同化物分配的生物地球化学模型和N、P富集下植物光合固C耦合模型研究,并应用同位素标记和分子生物学技术,从生态系统角度综合探讨N输入下植物光合固C的分配和转化特征。