Nitrogen and Phosphorus Limitation of Phytoplankton Growth in New Zealand Lakes: Implications for Eutrophication Control

We examine macronutrient limitation in New Zealand (NZ) lakes where, contrary to the phosphorus (P) only control paradigm, nitrogen (N) control is widely adopted to alleviate eutrophication. A review of published results of nutrient enrichment experiments showed that N more frequently limited lake productivity than P; however, stoichiometric analysis of a sample of 121 NZ lakes indicates that the majority (52.9%) of lakes have a mean ratio of total nitrogen (TN) to total phosphorus (TP) (by mass) indicative of potential P-limitation (>15:1), whereas only 14.0% of lakes have mean TN:TP indicative of potential N-limitation (<7:1). Comparison of TN, TP, and chlorophyll a data between 121 NZ lakes and 689 lakes in 15 European Union (EU) countries suggests that at the national scale, N has a greater role in determining lake productivity in NZ than in the EU. TN:TP is significantly lower in NZ lakes across all trophic states, a difference that is driven primarily by significantly lower in-lake TN concentrations at low trophic states and significantly higher TP concentrations at higher trophic states. The form of the TN:TP relationship differs between NZ and the EU countries, suggesting that lake nutrient sources and/or loss mechanisms differ between the two regions. Dual control of N and P should be the status quo for lacustrine eutrophication control in New Zealand and more effort is needed to reduce P inputs.     

Nutrient and Temperature Limitation of Bacterioplankton Growth in Temperate Lakes

Limitation of bacterioplankton production by nutrients and temperature was investigated in eight temperate lakes in summer. Six of the lakes were resampled in autumn. The lakes differ in nutrient content, water color, and concentration of dissolved organic carbon. Nutrients (phosphorus, nitrogen, and organic carbon) were added alone and in all possible combinations to filtered lake water inoculated with bacteria from the lake. After incubation for 36–40 h at in situ temperatures (ranging from 7 to 20°C), the response in bacterioplankton production was determined. The effect of increased temperature on bacterioplankton growth was also tested. Bacterioplankton production was often limited by phosphorus alone, organic carbon alone, or the two in combination. Phosphorus limitation of bacterioplankton production was more common in the summer, whereas limitation by organic carbon was more frequently observed in the autumn. There was a close balance between limitation by phosphorus and organic carbon in the epilimnion in the summer. In the hypolimnion in the summer, bacterioplankton growth was primarily phosphorus-limited. The effect of phosphorus additions decreased with increasing phosphorus concentrations in the lakes. However, there were no correlations between the effect of added organic carbon and water color, dissolved organic carbon concentration, or phosphorus concentration. When temperature was low (in the hypolimnion in the summer, and throughout the water column in the autumn) temperature also limited bacterioplankton production. Thus, temperature and inorganic nutrients or organic compounds can limit bacterioplankton growth both alone and simultaneously. However, at low temperatures, temperature is the most important factor influencing bacterioplankton growth.     

Nitrogen and Phosphorus Limitation over Long-Term Ecosystem Development in Terrestrial Ecosystems

Nutrient limitation to net primary production (NPP) displays a diversity of patterns as ecosystems develop over a range of timescales. For example, some ecosystems transition from N limitation on young soils to P limitation on geologically old soils, whereas others appear to remain N limited. Under what conditions should N limitation and P limitation prevail? When do transitions between N and P limitation occur? We analyzed transient dynamics of multiple timescales in an ecosystem model to investigate these questions. Post-disturbance dynamics in our model are controlled by a cascade of rates, from plant uptake (very fast) to litter turnover (fast) to plant mortality (intermediate) to plant-unavailable nutrient loss (slow) to weathering (very slow). Young ecosystems are N limited when symbiotic N fixation (SNF) is constrained and P weathering inputs are high relative to atmospheric N deposition and plant N:P demand, but P limited under opposite conditions. In the absence of SNF, N limitation is likely to worsen through succession (decades to centuries) because P is mineralized faster than N. Over long timescales (centuries and longer) this preferential P mineralization increases the N:P ratio of soil organic matter, leading to greater losses of plant-unavailable N versus P relative to plant N:P demand. These loss dynamics favor N limitation on older soils despite the rising organic matter N:P ratio. However, weathering depletion favors P limitation on older soils when continual P inputs (e.g., dust deposition) are low, so nutrient limitation at the terminal equilibrium depends on the balance of these input and loss effects. If NPP switches from N to P limitation over long time periods, the transition time depends most strongly on the P weathering rate. At all timescales SNF has the capacity to overcome N limitation, so nutrient limitation depends critically on limits to SNF.     

Estimating Summer Nutrient Concentrations in Northeastern Lakes from SPARROW Load Predictions and Modeled Lake Depth and Volume

Global nutrient cycles have been altered by the use of fossil fuels and fertilizers resulting in increases in nutrient loads to aquatic systems. In the United States, excess nutrients have been repeatedly reported as the primary cause of lake water quality impairments. Setting nutrient criteria that are protective of a lakes ecological condition is one common solution; however, the data required to do this are not always easily available. A useful solution for this is to combine available field data (i.e., The United States Environmental Protection Agency (USEPA) National Lake Assessment (NLA)) with average annual nutrient load models (i.e., USGS SPARROW model) to estimate summer concentrations across a large number of lakes. In this paper we use this combined approach and compare the observed total nitrogen (TN) and total phosphorus (TN) concentrations in Northeastern lakes from the 2007 National Lake Assessment to those predicted by the Northeast SPARROW model. We successfully adjusted the SPARROW predictions to the NLA observations with the use of Vollenweider equations, simple input-output models that predict nutrient concentrations in lakes based on nutrient loads and hydraulic residence time. This allows us to better predict summer concentrations of TN and TP in Northeastern lakes and ponds. On average we improved our predicted concentrations of TN and TP with Vollenweider models by 18.7% for nitrogen and 19.0% for phosphorus. These improved predictions are being used in other studies to model ecosystem services (e.g., aesthetics) and dis-services (e.g. cyanobacterial blooms) for ~18,000 lakes in the Northeastern United States.     

Diversity and Seasonal Dynamics of Actinobacteria Populations in Four Lakes in Northeastern Germany

The phylogenetic diversity and seasonal dynamics of freshwater Actinobacteria populations in four limnologically different lakes of the Mecklenburg-Brandenburg Lake District (northeastern Germany) were investigated. Fluorescence in situ hybridization was used to determine the seasonal abundances and dynamics of total Actinobacteria (probe HGC69a) and the three actinobacterial subclusters acI, acI-A, and acI-B (probes AcI-852, AcI-840-1, and AcI-840-2). Seasonal means of total Actinobacteria abundances in the epilimnia of the lakes varied from 13 to 36%, with maximum values of 30 to 58%, of all DAPI (4′,6′-diamidino-2-phenylindole)-stained cells. Around 80% of total Actinobacteria belonged to the acI cluster. The two subclusters acI-A and acI-B accounted for 60 to 91% of the acI cluster and showed seasonal means of 49% (acI-B) and 23% (acI-A) in relation to the acI cluster. Total Actinobacteria and members of the clusters acI and acI-B showed distinct seasonal changes in their absolute abundances, with maxima in late spring and fall/winter. In eight clone libraries constructed from the lakes, a total of 76 actinobacterial 16S rRNA gene sequences were identified from a total of 177 clones. The majority of the Actinobacteria sequences belonged to the acI and acIV cluster. Several new clusters and subclusters were found (acSTL, scB1-4, and acIVA-D). The majority of all obtained 16S rRNA gene sequences are distinct from those of already-cultured freshwater Actinobacteria.     

Absorption of Nitrogen, Phosphorus, and Potassium from Nutrient Sprays by Leaves

Barley, Brussels sprout, French bean, tomato, and sugar-beetplants grown in soil in pots and sprayed, usually daily, forseveral weeks, with nutrient solutions containing nitrogen,phosphorus, potassium, and a spreader, with precautions to preventthe spray solution falling on the soil, had higher nutrientcontents and dry weights than control plants sprayed with waterand spreader only. Increase in nutrient content occurred withhigh or low levels of nutrient supply to the roots and was approximatelyproportional to the concentration of spray and to the frequencyof spraying. The nitrogen content of sugar-beet plants was increased equallyby spraying with solutions supplying ammonium sulphate, calciumnitrate, or urea in equivalent concentrations. Nutrient uptake from solutions sprayed on leaves influenceduptake by the roots so that the additional amounts of nutrientcontained in sprayed plants may be greater or smaller than theamount absorbed from the spray by the leaves.     

N、P营养盐对塔玛亚历山大藻(Alexandrium tamarense)生长的影响

模拟自然海水营养盐浓度状况,在N、P浓度分别为10-500μg L-1 N和0.74-74μg L-1 P时,研究N、P双因子限制(N、P浓度同时降低,N:P固定为15:1)及单因子限制(保持N或P为最高浓度,只降低一种营养盐浓度)对有毒赤潮藻塔玛亚历山大藻(Alexandrium tamarense)生长的影响。结果表明,塔玛亚历山大藻细胞能较快进入对数生长期,但N、P双因子限制能明显影响其生长,在N、P浓度分别低于100μg L-1 N和15μg L-1 P时,细胞密度无明显增长;而N或P分别受限时,生长态势明显优于N、P同时受到限制的试验组,而且N、P单因子中度限制对生长影响较小。结果说明塔玛亚历山大藻对单因子营养元素限制较强的适应能力,可使其在常常出现单营养因子限制的自然水体中维持一定生长速率和细胞密度,并有助于滤食该藻的贝类体内麻痹性贝类毒素的积累。     

Controls of Benthic Nitrogen Fixation and Primary Production from Nutrient Enrichment of Oligotrophic, Arctic Lakes

We examined controls of benthic dinitrogen (N₂) fixation and primary production in oligotrophic lakes in Arctic Alaska, Toolik Field Station (Arctic Long-Term Ecological Research Site). Primary production in many oligotrophic lakes is limited by nitrogen (N), and benthic processes are important for whole-lake function. Oligotrophic lakes are increasingly susceptible to low-level, non-point source nutrient inputs, yet the effects on benthic processes are not well understood. This study examines the results from a whole-lake fertilization experiment in which N and P were added at a relatively low level (4 times natural loading) in Redfield ratio to a shallow (3 m) and a deep (20 m) oligotrophic lake. The two lakes showed similar responses to fertilization: benthic primary production and respiration (each 50–150 mg C m^?2 day^?1) remained the same, and benthic N₂ fixation declined by a factor of three- to fourfold by the second year of treatment (from ~0.35 to 0.1 mg N m^?2 day^?1). This showed that the response of benthic N₂ fixation was de-coupled from the nutrient limitation status of benthic primary producers and raised questions about the mechanisms, which were examined in separate laboratory experiments. Bioassay experiments in intact cores also showed no response of benthic primary production to added N and P, but contrasted with the whole-lake experiment in that N₂ fixation did not respond to added N, either alone or in conjunction with P. This inconsistency was likely a result of nitrogenase activity of existing N₂ fixers during the relative short duration (9 days) of the bioassay experiment. N₂ fixation showed a positive saturating response when light was increased in the laboratory, but was not statistically related to ambient light level in the field, leading us to conclude that light limitation of the benthos from increasing water-column production was not important. Thus, increased N availability in the sediments through direct uptake likely caused a reduction in N₂ fixation. These results show the capacity of the benthos in oligotrophic systems to buffer the whole-system response to nutrient addition by the apparent ability for significant nutrient uptake and the rapid decline in N₂ fixation in response to added nutrients. Reduced benthic N₂ fixation may be an early indicator of a eutrophication response of lakes which precedes the transition from benthic to water-column-dominated systems.     

Seasonal Trends in Nitrogen Status of Antarctic Lichens

The thallus nitrogen (N) concentration of two dominant macrolichensof continental Antarctica (Usnea sphacelata and Umbilicariadecussata) was estimated each month for 1 year on a low roundedknoll on Clark Peninsula, Windmill Islands, Wilkes Land, Antarctica.Thallus N was significantly higher in Umbilicaria decussatathan in Usnea sphacelata and varied according to site. Duringthe winter months, when the lichens were metabolically inactive,thalli gradually accumulated N. At the onset of warmer conditions,thallus N content fell dramatically in both species, with thetiming of the decline being related to microclimatic conditions.The strongly seasonal pattern of metabolic activity in thesespecies is reflected in their nitrogen relations. Copyright2000 Annals of Botany Company Lichen, nitrogen, Antarctica, season, Usnea sphacelata, Umbilicaria decussata     

Relationship between the Relative Limitation and Resorption Efficiency of Nitrogen vs Phosphorus in Woody Plants

Most previous studies have ascribed variations in the resorption of a certain plant nutrient to its corresponding environmental availability or level in tissues, regardless of the other nutrients’ status. However, given that plant growth relies on both sufficient and balanced nutrient supply, the nutrient resorption process should not only be related to the absolute nutrient status, but also be regulated by the relative limitation of the nutrient. Here, based on a global woody-plants dataset from literature, we test the hypothesis that plants resorb proportionately more nitrogen (or phosphorus) when they are nitrogen (or phosphorus) limited, or similar proportions of nitrogen (N) and phosphorus (P) when co-limited by both nutrients (the relative resorption hypothesis). Using the N:P ratio in green foliage as an indicator of nutrient limitation, we found an inverse relationship between the difference in the proportionate resorption of N vs P and this foliar N:P ratio, consistent across species, growth-forms, and vegetation-types globally. Moreover, according to the relative resorption hypothesis, communities with higher/lower foliar N:P (more likely P/N limited) tend to produce litter with disproportionately higher/lower N:P, causing a worsening status of P/N availability; this positive feedback may somehow be counteracted by several negative-feedback mechanisms. Compared to N, P generally shows higher variability in resorption efficiency (proportion resorbed), and higher resorption sensitivity to nutrient availability, implying that the resorption of P seems more important for plant nutrient conservation and N:P stoichiometry. Our findings elucidate the nutrient limitation effects on resorption efficiency in woody plants at the global scale, and thus can improve the understanding of nutrient resorption process in plants. This study also suggests the importance of the foliar N:P ratio as a key parameter for biogeochemical modeling, and the relative resorption hypothesis used to deduce the critical (optimal) N:P ratio for a specific plant community.     

Serum BDNF Concentrations Show Strong Seasonal Variation and Correlations with the Amount of Ambient Sunlight

Earlier findings show seasonality in processes and behaviors such as brain plasticity and depression that in part are regulated by Brain-Derived Neurotrophic Factor (BDNF). Based on this we investigated seasonal variation in serum BDNF concentrations in 2,851 persons who took part in the Netherlands Study of Depression and Anxiety (NESDA). Analyses by month of sampling (monthly n’s >196) showed pronounced seasonal variation in serum BDNF concentrations (P<.0001) with increasing concentrations in the spring-summer period (standardized regression weight (ß) = 0.19, P<.0001) and decreasing concentrations in the autumn-winter period (ß = −0.17, P<.0001). Effect sizes [Cohen’s d] ranged from 0.27 to 0.66 for monthly significant differences. We found similar seasonal variation for both sexes and for persons with a DSM-IV depression diagnosis and healthy control subjects. In explorative analyses we found that the number of sunshine hours (a major trigger to entrain seasonality) in the week of blood withdrawal and the 10 weeks prior to this event positively correlated with serum BDNF concentrations (Pearson’s correlation coefficients ranged: 0.05 – 0.18) and this could partly explain the observed monthly variation. These results provide strong evidence that serum BDNF concentrations systematically vary over the year. This finding is important for our understanding of those factors that regulate BDNF expression and may provide novel avenues to understand seasonal dependent changes in behavior and illness such as depression. Finally, the findings reported here should be taken into account when designing and interpreting studies on BDNF.     

Effects of Phosphorus Removal on the Maximal Algal Growth in Bioassay Experiments with Water from Four Dutch Lakes

This paper reports the effects of phosphorus removal at three sewage wastewater treatment plants on the state of eutrophication of four shallow lakes in the south-eastern part of the Rijnland Waterboard area during the years 1980–1982. With chemical analyses and bioassay experiments using the natural phytoplankton population no significant lowering could be detected of respectively the phosphate concentration and the maximal algal growth potential. All lakes proved to be principally nitrogen limited except the Reeuwijk Lakes, which showed clearly, after a primary nitrogen limitation, a secondary phosphorus limitation. Therefore the main attention with respect to phosphorus reduction should be concentrated on the Reeuwijk Lakes in the first place. For the other lakes in the investigated area phosphorus removal will, when it is the only measure taken, presumably not lead at short notice to a decrease of the algal biomass.     

Growth and Nitrogen Status of Soilless Tomato Plants Following Nitrate Withdrawal from the Nutrient Solution

The effects of withdrawing nitrogen (N) from the nutrient solutionof adult tomato plants growing in rockwool in a greenhouse wereinvestigated over a 6 week period during fruit production. Thetreatment reduced total plant growth after a lag period of about2 weeks. The commercial fruit yield after 6 weeks of N deprivationwas 7.7 kg m^-2compared to 9.3 kg m^-2in control plants. Duringthe experiment, growth of the -N plants was fuelled by N reservescontained in both the substrate (rockwool) and in plant organs.The nitrogen budget calculated for -N plants showed that onlya small amount of organic-N was readily available for internalcycling from organs such as stems. It served mainly to feedgrowing fruits which were the main sinks in the plant. The studyalso established that stores of nitrate-N were fully depletedbut it took 45 d for the -N plants to metabolize completelytheir nitrate reserves. This indicates that internal nitrateis not a readily-accessible store of labile N. An estimationof the critical N concentration (%N_c) in the aerial dry matterwas made from the data. Thus, for a crop yielding about 9.9tons DM ha^-1, %N_cwas close to 2.5%. This result is discussedin light of existing models that describe the ontogenic declinein %N_cin dry biomass of C₃plants. The study indicates that thecurrent regime of N fertilization practised in soilless culturesnot only leads to ineffective nitrogen use but also to largelosses of N to the environment; N concentrations should be decreasedin feeding recipes. The use of N-free nutrient solutions priorto the termination of plant culture may also be a means of limitingthe loss of eutrophying elements, such as nitrate, to the environment.Copyright 2001 Annals of Botany Company Lycopersicon esculentum, tomato, organ dry biomass, critical nitrogen concentration, compartment, rockwool, nitrate interruption, distribution, reserves     

Nutrient Limitation in Three Lowland Tropical Forests in Southern China Receiving High Nitrogen Deposition: Insights from Fine Root Responses to Nutrient Additions

Elevated nitrogen (N) deposition to tropical forests may accelerate ecosystem phosphorus (P) limitation. This study examined responses of fine root biomass, nutrient concentrations, and acid phosphatase activity (APA) of bulk soil to five years of N and P additions in one old-growth and two younger lowland tropical forests in southern China. The old-growth forest had higher N capital than the two younger forests from long-term N accumulation. From February 2007 to July 2012, four experimental treatments were established at the following levels: Control, N-addition (150 kg N ha^–1 yr^–1), P-addition (150 kg P ha^–1 yr^–1) and N+P-addition (150 kg N ha^–1 yr^–1 plus 150 kg P ha^–1 yr^–1). We hypothesized that fine root growth in the N-rich old-growth forest would be limited by P availability, and in the two younger forests would primarily respond to N additions due to large plant N demand. Results showed that five years of N addition significantly decreased live fine root biomass only in the old-growth forest (by 31%), but significantly elevated dead fine root biomass in all the three forests (by 64% to 101%), causing decreased live fine root proportion in the old-growth and the pine forests. P addition significantly increased live fine root biomass in all three forests (by 20% to 76%). The combined N and P treatment significantly increased live fine root biomass in the two younger forests but not in the old-growth forest. These results suggest that fine root growth in all three study forests appeared to be P-limited. This was further confirmed by current status of fine root N:P ratios, APA in bulk soil, and their responses to N and P treatments. Moreover, N addition significantly increased APA only in the old-growth forest, consistent with the conclusion that the old-growth forest was more P-limited than the younger forests.     

Seasonal Predictability of the East Atlantic Pattern from Sea Surface Temperatures

This study analyzes the influence of sea surface temperatures (SSTs) on the second mode of atmospheric variability in the north Atlantic/European sector, namely the East-Atlantic (EA) pattern, for the period 1950–2012. For this purpose, lead-lag relationships between SSTs and the EA pattern, ranging from 0 to 3 seasons, were assessed. As a main result, anomalies of the EA pattern in boreal summer and autumn are significantly related to SST anomalies in the Indo-Pacific Ocean during the preceding seasons. A statistical forecasting scheme based on multiple linear regression was used to hindcast the EA-anomalies with a lead-time of 1 to 2 months. The results of a one-year-out cross-validation approach indicate that the phases of the EA in summer and autumn can be properly hindcast.     

Temporal Patterns in Bacterial Communities in Three Temperate Lakes of Different Trophic Status

Despite considerable attention in recent years, the composition and dynamics of lake bacterial communities over annual time scales are poorly understood. This study used automated ribosomal intergenic spacer analysis (ARISA) to explore the patterns of change in lake bacterial communities in three temperate lakes over 2 consecutive years. The study lakes included a humic lake, an oligotrophic lake, and a eutrophic lake, and the epilimnetic bacterial communities were sampled every 2 weeks. The patterns of change in bacterial communities indicated that seasonal forces were important in structuring the behavior of the bacterial communities in each lake. All three lakes had relatively stable community composition in spring and fall, but summer changes were dramatic. Summertime variability was often characterized by recurrent drops in bacterial diversity. Specific ARISA fragments derived from these lakes were not constant among lakes or from year to year, and those fragments that did recur in lakes in different years did not exhibit the same seasonal pattern of recurrence. Nonetheless, seasonal patterns observed in 2000 were fairly successful predictors of the rate of change in bacterial communities and in the degree of autocorrelation of bacterial communities in 2001. Thus, seasonal forces may be important structuring elements of these systems as a whole even if they are uncoupled from the dynamics of the individual system components.     

不同氮磷浓度对莱布新月藻生长与脂溶性物质含量的影响

以莱布新月藻Closterium leibleinii为试材,利用单因子分析研究不同氮、磷浓度培养的莱布新月藻生长及脂溶性物质含量变化,为其作为生物柴油开发提供基础资料。结果表明,莱布新月藻生长的最适氮浓度为5.30 mg·L^–1,其最大生长速率为3.675×10⁶个·L^–1·d^–1;最适磷浓度为0.02 mg·L^–1,最大生长速率为4.05×10⁵个·L^–1·d^–1。在最适氮磷浓度BG11培养下粗脂约占藻体干重的88.04%。莱布新月藻在适宜的氮磷浓度培养基中生长速率较快,脂溶性物质含量高,可作为一种较有潜力的植物开发生物柴油。     

Phytoplankton and Epiphyte Development and Their Shading Effect on Submerged Macrophytes in Lakes of Different Nutrient Status

The annual mean light intensity at the depth limit of the Littorella vegetation was 24–33% of the subsurface light intensity, despite large variations in each attenuation component (lake water, phytoplankton, and epiphytes). In oligotrophic, silicate-poor lakes, the light attenuation above the submerged vegetation was dominated by the water itself, which accounted for 65–72% of the total attenuation. Phytoplankton and epiphytes were equal in importance to each other. In oligotrophic, silicate-rich lakes and lakes receiving a nitrogen supply above background level, the epiphytes were more abundant, accounting for about 50% of the light attenuation. In one lake with a high nutrient supply, the epiphytes were responsible for 86% of the light attenuation. A new method of measuring the effect of shading by the epiphytic community on submerged macrophytes is presented. The light attenuation caused by the phytoplankton and the epiphytes was investigated and related to the depth distribution of the submerged angiosperm, Littorella uniflora. It is shown that the biomass of the epiphytes increased more than the biomass of the phytoplankton in response to an external or internal nutrient loading. Shading by epiphytes is of decisive importance for the depth distribution of Littorella at increasing nutrient supply.