Horizontal variation of biomass and C:N:P ratios of benthic algae in lakes

The horizontal variation of chlorophyll a (chl a) and C:N:P (carbon:nitrogen:phosphorus) ratio was estimated for benthic algal communities attached to living substrates (mussels and macrophytes) and to rocks and stones in three lakes of different trophy. Samples were taken in a nested hierarchical design with replicates separated by several cm, dm, 10 m, and km. The observed horizontal variation of chl a, C:N, and C:P ratios did not differ for horizontal scales, substrates, or lakes. Although the investigated lakes were quite unlike regarding nutrient status, light regime, or morphology, the patchiness was similar in all lakes. Moreover, patchiness was also similar on stones, macrophytes and mussels, although those substrates differed in longevity and surface structure. Similar patchiness regardless of scale, substrate, or sampled lake, implies the possibility of using an optimal sampling design calculated for one lake and substrate also in other lakes and on other substrates.     

Seasonal variability of N:P ratios in eutrophic lakes

The ratios of different forms of nitrogen to phosphorus (particulate, total, total dissolved, and dissolved inorganic N : P) from four eutrophic lakes are presented to assess their variability during two consecutive growing seasons. Particulate and total N : P ratios showed the lowest amplitude of fluctuations, while the total inorganic N : P ratios showed the highest. All N : P forms demonstrated substantial variations and seasonal pattern over the growing period April-October. It was the short-lasting spring minima of the N : P ratios, ranging from less than 1: 1 to 6 : 1, which triggered the onset of nitrogen-fixing cyanophyte blooms. Seasonal mean values were as high as 20: 1 to 30: 1 and misleading in assessing nitrogen limitation. The nitrogen fixation rapidly restored the N : P values to normal levels (TN: TP of 15: 1 and over).     

Benthic algal response to N and P enrichment along a pH gradient

Nutrient enrichment and its effect on benthic algal growth, community composition, and average cell size was assessed across two sites of differing pH within a single habitat. Nutrients were added using in situ substrata, which released either N, P, or no additional nutrients (controls) at each site for 21 days. Upon collection, chlorophyll and biovolume standing stocks of the attached algal microflora were measured. Chlorophyll concentration was different among all treatments, accumulating greatest on P, followed by N, and the least on C substrata (P < 0.001) and was highest at site-2 (P < 0.001), while total algal biovolume was highest on P compared to both N and C substrata (P < 0.05) and did not vary between sites. Increased growth on P substrata was due to the enhanced biovolume of filamentous green algae, although the affected taxa varied between sites. Biovolume to cell density ratios (as a measure of average cell size) were highest on P substrata over both N-enriched and control substrata (P < 0.05) and this pattern was similar between sites. Progression towards a community composed of larger cells following P enrichment observed along this pH gradient, seems to be related to the dominance of larger celled filamentous green algae. Thus, nutrients exhibited greater control on benthic algal growth than did changes in hydrogen ion concentration.Contribution number 581, Great Lakes Environmental Research LaboratoryContribution number 581, Great Lakes Environmental Research Laboratory     

Combined effects of nitrogen enrichment, sulphur pollution and climate change on fen meadow vegetation N:P stoichiometry and biomass

Nitrogen (N) and sulphur (S) deposition, as well as altered soil moisture dynamics due to climate change can have large effects on fen meadow biogeochemistry and vegetation. Their combined effects may differ strongly from their separate effects, since each process affects different nutrients through different mechanisms. However, the impacts of these environmental problems are rarely studied in combination. We therefore investigated the separate and interactive effects of current levels of N- and S-deposition and changes in soil moisture dynamics on fen meadow vegetation. We focused on vegetation biomass and N:P stoichiometry, including access to soil P through root surface phosphatase activity, in a 3-year factorial addition experiment in an N-limited rich fen meadow in the Biebrza valley in Poland. We applied 29.5 kg N ha^?1 year^?1 and 32.1 kg S ha^?1 year^?1, which correspond to current deposition levels in Western Europe. Changes in soil moisture dynamics due to climate change were mimicked by amplified drying of the soil in summer. This level of N-deposition had limited effects on plant biomass production in this rich fen, despite low foliar N:P ratios that suggest N limitation. This level of S-deposition, however, resulted in decreased vegetation P-uptake and biomass. We also showed that increased summer drought resulted in considerable increases in vegetation biomass. We found no interactive effects on vegetation biomass or N:P stoichiometry, possibly as a result of the limited main effects of the separate processes.     

Nutrient recycling and shifts in N:P ratio by different zooplankton structures in a South Andes lake

In South Andes lakes, zooplankton succession is characterizedby a change in the dominance from the calanoid copepod Boeckellagracilipes in winter and spring, to the cladoceran Bosmina longirostrisin midsummer, and the rotifer Polyarthra vulgaris in late summer.We performed three series of field experiments at differenttimes (late spring, summer and late summer) to examine the roleof zooplankton constitution in the released nitrogen:phosphorus(N:P) ratio. We observed that changes in the zooplankton constitutionover the annual cycle may change the nutrient supply ratio.In South Andes lakes, Boeckella gracilipes would decrease theP limitation, lowering the N:P ratio, whereas Bosmina longirostristends to increase the N:P ratio and therefore increase the Plimitation during summer.     

Relationships among nitrogen and total phosphorus,algal biomass and zooplankton density in the central Amazonia lakes

The relationship among concentrations of total nitrogen (TN), total phosphorus (TP), algal biomass (Chl) and the density and size of individuals of the zooplankton community were studied for the dry season (November 1999–January 2000) at 20 lakes of the Central Amazonia. The study was conducted along a productivity gradient to identify the existence of resource or predator-dependent patterns on the primary producers of the trophic web. A strong positive relationship was observed between the log Chl and TN (r ² = 0.88, P = 0.000) and to log Chl and log TP (r ² = 0.85, P = 0.000) in a simple linear regression. However, when both variables were running together in a multiple regression, TN alone explained every variation of algal biomass (r ² = 0.89, P _TN = 0.022, P _TP = 0.233). The total density of the zooplankton showed a positive correlation with log Chl (r ² = 0.53, P = 0.000) and the large zooplankton (>0.5 mm) was found to be a more positive function of the phytoplankton (r ² = 0.65) than the density of the small ones (<0.5 mm, r ² = 0.44). Results show that complex food web interactions could be responsible for patterns in tropical systems. We contend that Chl variation in tropical lake systems is controlled by TN and TP, but the predictor power of the TN increase the fit of the model in analysis and can be use alone to access the variability in algae biomass to Amazonian tropical lakes. We also agree that the density of large zooplankton individuals is regulated by the biomass of primary producers. Hence we concluded that the resource-dependent hypothesis is supported in these systems. Handling editor: J. Padisak     

Phylogenetic ecology of foliar N and P concentrations and N:P ratios across mediterranean‐type ecosystems

Aim Interpretations of global‐scale patterns in foliar N and P concentrations and N:P ratios across climatic gradients and biomes often ignore complications imposed by taxonomic and phylogenetic structure in the nutrient concentrations of the constituent taxa. We analysed foliar N and P concentrations and N:P ratios of species from similar climate zones, but with contrasting soil fertility, to determine the relative importance of phylogeny and geographic region in driving foliar nutrient concentrations. Location Mediterranean climate regions. Methods Mean foliar N and P concentrations of 564 species from five mediterranean climate regions were compiled. Regional comparisons of foliar N and P concentrations and N:P ratios were performed using a phylogenetically independent contrasts (PIC) procedure. We also evaluated phylogenetic structuring in these variables, determining for each clade whether (1) the mean trait value and (2) the variance in trait values deviate significantly from chance expectation. Results Foliar N and P concentrations were found to vary between regions, a result confirmed using PIC. Tests of phylogenetic structure identified lineages having unusually high or low N and P concentrations, these being largely consistent amongst two of the regions. There was a general pattern of conservatism in foliar N and P concentrations and N:P ratios with localized instances of overdispersion. Main conclusions Our findings identify soil fertility as a strong environmental filter which has led to the dominance of low‐nutrient adapted lineages in the South African Cape and Western Australia. There is also a pronounced clade‐specific ‘stoichiometric homeostasis’ in foliar N and P concentrations and N:P ratios and this strong phylogenetic conservatism, together with the narrow range of foliar concentrations, is an evolutionary outcome of factors associated with developmental and physiological nutrient requirements specific to each lineage. Our results reinforce the need to include phylogenetic considerations in discussions of the biological stoichiometry of plants if we are to expand our current knowledge of foliar N and P concentrations and N:P patterns from an ecological to a biogeographic scale.     

Response of zooplankton to nutrient enrichment and fish in shallow lakes: a pan-European mesocosm experiment

1. Responses of zooplankton to nutrient enrichment and fish predation were studied in 1998 and 1999 by carrying out parallel mesocosm experiments in six lakes across Europe. 2. Zooplankton community structure, biomass and responses to nutrient and fish manipulation showed geographical and year‐to‐year differences. Fish had a greater influence than nutrients in regulating zooplankton biomass and especially the relative abundances of different functional groups of zooplankton. When fish reduced the biomass of large crustaceans, there was a complementary increase in the biomasses of smaller crustacean species and rotifers. 3. High abundance of submerged macrophytes provided refuge for zooplankton against fish predation but this refuge effect differed notably in magnitude among sites. 4. Large crustacean grazers (Daphnia, Diaphanosoma, Sida and Simocephalus) were crucial in controlling algal biomass, while smaller crustacean grazers and rotifers were of minor importance. Large grazers were able to control phytoplankton biomass even under hypereutrophic conditions (up to 1600 μg TP L^?1) when grazer biomass was high (>80–90 μg dry mass L^?1) or accounted for >30% of the grazer community. 5. The littoral zooplankton community was less resistant to change following nutrient enrichment in southern Spain, at high temperatures (close to 30 °C), than at lower temperatures (17–23 °C) characterising the other sites. This lower resistance was because of a greater importance of nutrients than zooplankton in controlling algal biomass. 6. Apart from the reduced role of large crustacean grazers at the lowest latitude, no consistent geographical patterns were observed in the responses of zooplankton communities to nutrient and fish manipulation.     

Effects of nutrients and fish on periphyton and plant biomass across a European latitudinal gradient

Replicated, factorial mesocosm experiments were conducted across Europe to study the effects of nutrient enrichment and fish density on macrophytes and on periphyton chlorophyll a (chl-a) with regard to latitude. Periphyton chl-a densities and plant decline were significantly related to nutrient loading in all countries. Fish effects were significant in a few sites only, mostly because of their contribution to the nutrient pool. A saturation-response type curve in periphyton chl-a with nutrients was found, and northern lakes achieved higher densities than southern lakes. Nutrient concentration and phytoplankton chl-a necessary for a 50% plant reduction followed a latitudinal gradient. Total phosphorus values for 50% plant disappearance were similar from Sweden (0.27 mg L⁻¹) to northern Spain (0.35 mg L⁻¹), but with a sharp increase in southern Spain (0.9 mg L⁻¹). Planktonic chl-a values for 50% plant reduction increased monotonically from Sweden (30 μg L⁻¹) to València (150 μg L⁻¹). Longer plant growing-season, higher light intensities and temperature, and strong water-level fluctuations characteristic of southern latitudes can lead to greater persistence of macrophyte biomass at higher turbidities and nutrient concentration than in northern lakes. Results support the evidence that latitudinal differences in the functioning of shallow lakes should be considered in lake management and conservation policies.     

Effects of available P and N:P ratios on non-symbiotic dinitrogen fixation in tallgrass prairie soils

Summary Prescribed burning is a major control over element cycles in Tallgrass prairie (Eastern Kansas, USA). In this paper we report potential effects of fire on nonsymbiotic nitrogen fixation. Fire resulted in additions of available P in ash, which may stimulate nitrogen fixation by terrestrial cyanobacteria. Cyanobacterial nitrogenase activity and biomass responded positively to additions of ash or P in laboratory assays using soil. Further assays in soil showed that cyanobacteria responded to changes in available N:available P ratio (aN:P) across a range of concentrations. Nitrogen fixation rate could be related empirically to aN:P via a log-linear relationship. Extrapolation of laboratory results to the field yielded a maximal estimate of 21 kg N ha^-1 y^-1. Results support arguments from the marine and terrestrial literature that P availability is central to regulation of ecosystem N budgets.     

Plant community response to nitrogen and phosphorus enrichment varies across an alpine tundra moisture gradient

Background: The extent to which nutrient availability influences plant community composition and dynamics has been a focus of ecological enquiry for decades.

Aims: Results from a long-term nitrogen (N) and phosphorus (P) addition experiment in alpine tundra were used to evaluate the importance of the two nutrients in structuring plant communities in three communities that differed in their snow cover amounts and duration and soil moisture characteristics.

Methods: A factorial N and P experiment was established in three meadows differing in initial vegetation composition and soil moisture. Plant and soil characteristics were measured after 20 years, and the dissimilarity among meadows and treatments were measured using permutational analysis of variance.

Results: Plant species richness declined uniformly across the three meadow types and in response to N and N + P additions, while both evenness and the Shannon diversity index finding indicated that nutrient additions had the highest impact on moister habitats. Overall, N impacts overshadowed changes attributed to P additions, and the N and N + P plots in wet meadow sites were the least diverse and scored the lowest dissimilarity averages among treatments. Dissimilarity estimates indicated that the control and P plots in the dry meadow community were more distinct in composition than all other plots, and especially those in the moist or wet meadows. Above-ground biomass of grasses and sedges (graminoids) increased with N additions while forbs appeared to show responses dictated in part by the graminoid responses. The most abundant grass species of moist and wet meadow, Deschampsia cespitosa, dominated N and N + P plots of the wet sites, but did not show a N response in moist areas in spite of its general abundance in moist meadow. Competition from other plant species in the moist areas likely diminished the D. cespitosa response and contributed to the resilience of the community to nutrient enrichment.

Conclusions: Initial community composition, as influenced by the specific moisture regime, appears to control the extent to which changes in nutrient resources can alter plant community structure. Long-term fertilization tends to support most but not all findings obtained from shorter-termed efforts, and wet meadows exhibit the largest changes in plant species numbers and composition when chronically enriched with N.     

Effects of fire history and N and P fertilization on seedling biomass,Specific Leaf Area,and root:shoot ratios in a South African savannah

Fire exerts clear direct effects on savannah vegetation dynamics, but the indirect effects of fire, for example via changes in soil fertility, are less clear. In fire-dominated nutrient-poor savannahs, there is little evidence that fire affects plant-available nutrients, but plant performance on soils with contrasting fire histories has not been investigated. A bioassay experiment was conducted in Kruger National Park, South Africa, to test for effects of long-term fire manipulation on the ability of soils to influence tree growth. Combretum hereroense, a common savannah species, was used as a phytometer species in an experiment that combined fire history with N and P fertilization treatments. The results suggested no effect of fire history on a number of seedling traits after seven months of growth. Root mass and shoot:root ratios responded to P addition, with seedlings exhibiting lower allocation to root tissue under elevated P, particularly in the presence of N, suggesting some N and P co-limitation. Overall, it is concluded that there is little evidence that fire degrades or alters soil chemistry in a way that is likely to influence vegetation structure and dynamics in nutrient-poor savannah ecosystems.     

Composition,size, and biomass of zooplankton in large productive Florida lakes

Crustacean zooplankton data were compiled from long-term observational studies at seven large shallow Florida lakes, to determine whether there are general characteristics in regard to species composition, body size, and biomass. In particular, we examined whether patterns in body size and species richness fit empirical models developed by Stanley Dodson. The lakes included range in size from 125 to 1730 km² and encompass mesotrophic to hyper-eutrophic conditions. We found that zooplankton biomass was strongly dominated by one species of calanoid copepod—Arctodiaptomus dorsalis. Large daphnids were absent, and Cladocera assemblages were dominated by small taxa such as Ceriodaphnia, Chydorus, and Eubosmina. The total number of species of pelagic cladocerans (8–12) was consistent with Dodson’s predictions based on lake area. The average size of crustacean zooplankton in Florida lakes is small in comparison with temperate communities. A. dorsalis is the smallest calanoid copepod in North America, and the mean length of Cladocera (0.6 mm) is consistent with Dodson’s results that size decreases from temperate to tropical zones. Total biomass of crustacean zooplankton was very low, ratios of zooplankton to phytoplankton biomass (0.01–0.1) are among the lowest reported in the literature, and the zooplankton displayed short-lasting early spring peaks in biomass. Cladocera were almost entirely absent in spring and summer. Factors known to occur in Florida lakes, which appear to explain these characteristics of biomass, include intense fish predation and high summer water temperature.     

Benthic macroinvertebrate biomass and wildfires: evidence for enrichment of boreal subarctic lakes

1 We quantified the effects of forest fire on littoral benthic macroinvertebrate biomass on a boreal subarctic plateau in Alberta, Canada. Water chemistry and benthos were collected from six lakes, 1 and 2 years following a 1995 fire which burned about 91% of their catchments (i.e. recently burnt lakes), and from four other lakes whose catchments burned between 1961 and 1985 (i.e. previously burnt lakes). Seven lakes whose catchments had not burned since at least 1957 served as reference systems.
2 Total benthic macroinvertebrate biomass and biomass of Chironomidae were 1.5‐ and 2‐fold (P<0.05) greater within recently burnt lakes than in reference systems, whereas the biomass of Oligochaeta (P=0.06) and Amphipoda (P=0.07) were marginally higher in burnt than reference lakes. Burnt lakes had greater colour (P<0.05) and marginally (P=0.06) higher concentrations of soluble reactive phosphorus than reference lakes.
3 Nutrient diffusing substrata deployed in a previously and a recently burnt lake indicated that littoral epilithic communities were co‐limited by the availability of phosphorus (P) and nitrogen (N), although co‐limitation was stronger in the previously burnt than the recently burnt lake. Epilithic chlorophyll a on natural stone surfaces in the recently burnt lake was also 3.5 times higher (P<0.05) than that from the previously burnt lake.
4 Among all 17 lakes, total benthic biomass and biomass of Chironomidae, Amphipoda and Nematoda, were significantly (P<0.05) or marginally (P=0.06) related to soluble reactive phosphorus (SRP) but not dissolved inorganic nitrogen or colour. These regressions explained between 11% and 64% of variation in benthic biomasses.
5 Linear regressions and second‐order polynomials explained 18% and 24% of the variation in concentrations of SRP and water colour with time since fire, respectively, and between 22% and 70% of variation in total biomass and biomass of the five dominant invertebrate groups. These analyses suggest that benthic biomasses continue to be elevated for about 15–20 years following fire before declining to pre‐disturbance levels.     

Effects of limiting nutrients and N:P ratios on the phytoplankton growth in a shallow hypertrophic reservoir

The purpose of this study was to evaluate the effects of limiting nutrients and the N:P ratios on the growth of phytoplankton (mainly cyanobacteria) in a shallow hypertrophic reservoir between November 2002 and December 2003. Nutrient enrichment bioassays (NEBs) were conducted, along with analyses of seasonal ambient nutrients and phytoplankton taxa, in the reservoir. The average DIN:TDP and TN:TP mass ratios in the ambient water were 90 (range: 17–187) and 34 (13–60), respectively, during the study period. The dissolved inorganic phosphorus showed seasonal variation, but less than that of inorganic nitrogen. The TN:TP ratios ranged from 13 to 46 (mean: 27 ± 6) during June–December when the cyanobacteria, Microcystis, dominated the phytoplankton composition. The NEBs showed that phytoplankton growth was mainly stimulated by the phosphorus (all of total 17 cases), rather than the nitrogen concentration (8 of 17 cases). The rapid growth rate of cyanobacteria was evident with TN:TP ratios less than 30. According to the results of the NEBs with different N concentrations (0.07, 0.7 and 3.5 mg l⁻¹), but the same N:P ratios and when the nitrogen concentration was highest, the cyanobacterial growth reached a maximum at N:P ratios <1. Overall, the response of cyanobacterial growth was a direct function of added phosphorus in the NEBs, and was greater with increased N concentrations. Thus, cyanobacterial blooms favored relatively low N:P ratios in this hypertrophic reservoir system. An erratum to this article is available at .     

无机氮磷添加对太湖来源浮游植物和附着生物生物量的影响术

为了探讨富营养对太湖来源水体浮游植物的影响,对来源于太湖梅梁湾的水体进行氮磷营养盐添加实验.实验共3个处理(T0、T1和T2),T0不进行营养盐添加,处理T1和T2添加无机氮磷,T1中总氮、总磷净增加分别为10和0.5 mg·L-1,T2添加的量为T1的2倍.结果表明:T0的浮游植物叶绿素a含量明显低于添加营养盐的2个处理,而添加营养盐的2个处理之间没有显著差异,说明氮磷营养盐的添加在一定程度上能够促进浮游植物的生长,但并非营养盐添加越高浮游植物生物量就越高;添加处理与对照中附着生物叶绿素a含量的比较结果与浮游植物生物量的相同;T2中浮游植物的生长可能受到除氮磷之外的因素的限制;T1和T2中均没有出现期望的蓝藻水华,表明蓝藻水华的形成不只是与充足的氮磷供应有关.     

The mineralization of nitrogen and phosphorus in organic materials of varying C:N and C:P ratios

Summary The nitrogen and phosphorus contents of organic materials as a factor in determining the trend of mineralization of these elements when plant materials are added to the soil, were studied in an incubation experiment over a period of 12 weeks.Nitrogen mineralization increased with decreasing C:N ratio. No nitrogen mineralization was recorded above C:N ratio of 16.1 and the critical ratio lay between this and 23.0, with the plant materials used.Initial phosphorus immobilization occurred when plant materials ranging in C:P ratio from 501 to 112 were decomposed in the soil. The release of the immobilized phosphorus as incubation progressed suggests that green manuring should be considered of long-term benefit as far as phosphorus is concerned.     

Effects of upstream lakes and nutrient limitation on periphytic biomass and nitrogen fixation in oligotrophic, subalpine streams

1. We conducted bioassays of nutrient limitation to understand how macronutrients and the position of streams relative to lakes control nitrogen (N₂) fixation and periphytic biomass in three oligotrophic Rocky Mountain catchments. We measured periphytic chlorophyll‐a (chl‐a) and nitrogen‐fixation responses to nitrogen (N) and phosphorus (P) additions using nutrient‐diffusing substrata at 19 stream study sites, located above and below lakes within the study catchments. 2. We found that periphytic chl‐a was significantly co‐limited by N and P at 13 of the 19 sites, with sole limitation by P observed at another four sites, and no nutrient response at the final two sites. On average, the addition of N, P and N + P stimulated chl‐a 35%, 114% and 700% above control values respectively. The addition of P alone stimulated nitrogen fixation by 2500% at five of the 19 sites. The addition of N, either with or without simultaneous P addition, suppressed nitrogen fixation by 73% at nine of the 19 sites. 3. Lake outlet streams were warmer and had higher dissolved organic carbon concentrations than inlet streams and those further upstream, but position relative to lakes did not affect chl‐a and nitrogen fixation in the absence of nutrient additions. Chl‐a response to nutrient additions did not change along the length of the study streams, but nitrogen fixation was suppressed more strongly by N, and stimulated more strongly by P, at lower altitude sites. The responses of chl‐a and nitrogen fixation to nutrients were not affected by location relative to lakes. Some variation in responses to nutrients could be explained by nitrate and/or total N concentration. 4. Periphytic chl‐a and nitrogen fixation were affected by nutrient supply, but responses to nutrients were independent of stream position in the landscape relative to lakes. Understanding interactions between nutrient supply, nitrogen fixation and chl‐a may help predict periphytic responses to future perturbations of oligotrophic streams, such as the deposition of atmospheric N.     

Effects of nutrient recycling by zooplankton and fish on phytoplankton communities

In laboratory experiments we tested the hypothesis that nutrients supplied by fish and zooplankton affect the structure and dynamics of phytoplankton communities. As expected from their body size differences, fish released nutrients at lower mass-specific rates than Daphnia. On average, these consumers released nutrients at similar N:P ratios, although the ratios released by Daphnia were more variable than those released by fish. Nutrient supply by both fish and Daphnia reduced species richness and diversity of phytoplankton communities and increased algal biomass and dominance. However, nutrient recycling by fish supported a more diverse phytoplankton community than nutrient recycling by Daphnia. We conclude that nutrient recycling by zooplankton and fish have different effects on phytoplankton community structure due to differences in the quality of nutrients released. Received: 21 December 1998 / Accepted: 31 May 1999     

Atmospheric nitrogen deposition has caused nitrogen enrichment and eutrophication of lakes in the northern hemisphere

We compiled chemical data and phytoplankton biomass (PB) data (chlorophyll a ) from unproductive lakes in 42 different regions in Europe and North America, and compared these data to inorganic nitrogen (N) deposition over these regions. We demonstrate that increased deposition of inorganic N over large areas of Europe and North America has caused elevated concentrations of inorganic N in lakes. In addition, the unproductive lakes in high N deposition areas had clearly higher PB relative to the total phosphorus (P) concentrations illustrating that the elevated inorganic N concentrations has resulted in eutrophication and increased biomass of phytoplankton. The eutrophication caused by inorganic N deposition indicates that PB yield in a majority of lakes in the northern hemisphere is (was) limited by N in their natural state. We, therefore, suggest that P limitation largely concerns lakes where the balance between N and P has been changed because of increased anthropogenic input of N.