The relationship between nutrient status and chemical composition of Peridinium cinctum during the bloom in Lake Kinneret

Changes in water chemistry and the major chemical and elementalconstituents of Peridinium cinctum have been followed duringthe 1976 growth season (February-June) in Lake Kinneret. Thedecline of the bloom was not associated with changes in anysingle physical or chemical parameter of the lake water, althoughthe ambient N:P ratio was higher before the peak of Peridinium.Carbon, nitrogen and chlorophyll in the cells were almost constantover the growth season, averaging 48, 4 and 0.9% dry weight,respectively. There was an initial drop in cell phosphorus from0.7% to 0.2% dry weight which then remained stable after April.During the bloom period the carbohydrate and lipid content ofPeridinium decreased, nucleic acid increased, whereas proteinshowed no definite trend. The results suggest that (a) Peridiniumis not severely nutrient limited during most of the growth season;and (b) criteria based upon cellular composition used to definenutrient status in other groups of algae, may not be directlyapplicable to armoured dinoflagellates.     

The role of phosphatases in the metabolism of Peridinium cinctum,from Lake Kinneret

The annual algal bloom (February–June) in Lake Kinneret consists almost entirely of the dinoflagellatePeridinium cinctum f.westii (Dinophyceae). To clarify the role of phosphatases in the alga, experiments were carried out using cells from culture or from the lake. In culture, as the external ambient orthophosphate (Pi) concentration decreased, alkaline phosphatase activity increased (and to some extent acid phosphatase activity, as well). Hot water extractable P decreased, although molybdate reactive phosphorus (MRP) appeared to be utilized in preference to the non-MRP component of this pool. Alkaline phosphatase inPeridinium collected from the lake as well as cells grown in culture under a high (3–6 mg l^–1) ambient Pi concentration in both continuous light and a 12:12 light-dark cycle, showed a diurnal fluctuation in activity. These results, together with previous observations suggest that the phosphatases inPeridinium are controlled by changes in intracellular phosphorus levels (other than the hot water extractable pool) and/or by other metabolic processes not directly involved in P nutrition.     

HOT WATER EXTRACTABLE PHOSPHORUS—AN INDICATOR OF NUTRITIONAL STATUS OF PERIDINIUM CINCTUM (DINOPHYCEAE) FROM LAKE KINNERET (ISRAEL)?1

The intracellular levels of hot water extractable and total phosphorus were determined in the dinoflagellate Peridinium cinctum. f. westii (Lemm.) Lef. for natural samples from the bloom in Lake Kinneret and from laboratory cultures. Amounts of phosphorus (P) in the hot water fraction, relative to total cellular phosphorus, were similar in lake Peridinium and in cells grown in high ambient orthophosphate (P_i) media (3–6 mg P · l^?1). The absolute amounts of hot water extractable P in natural cell and those cultured at lower P_i concentrations (0.02–0.05 mg P · 1^?1) were similar, although average P_i in lake water were 4 μg · l^?1. Under most growth conditions the hot water extract contained approximately equal amounts of molybdate reactive phosphorus (MRP) and non-MRP. Short chain (6–9 units) polyphosphates (mol wt 630–950) probably constituted the bulk of the non-MRP pool, which was hydrolysable by alkaline phosphatase and may serve as a precursor for a more permanent P store. Intracellular P levels and distribution were not directly dependent on external P_i concentrations but may be determined by the N:P atomic ratio or overall external ionic milieu. Peridinium grown in low ambient P_i released significant amounts of non-MRP compounds. In Lake Kinneret, for at least most of the bloom period, Peridinium does not appear to be limited by P supply.     

A mesocosm study of Phaeocystis globosa (Prymnesiophyceae) population dynamics: II. Significance for the microbial community

The impact of Phaeocystis globosa population decline on the microbial community was studied during a mesocosm experiment, with irradiance regime and inorganic N:P ratios (4, 16, and 44) as controlling factors. Heterotrophic bacterial activity was closely related to enhanced (viral) lysis rates of P. globosa cells and disintegration of the colonies. Up to 85% of the bacterial C demand could be supplied by P. globosa-specific cellular C release. The bacterial populations with high DNA content became dominant (>70% of total). The bacterial community showed a rapid shift in composition to take advantage of the changing conditions during the demise of the P. globosa bloom. Members of the Alphaproteobacteria and the Bacteroidetes group emerged directly upon bloom decay. Multidimensional scaling analysis in conjunction with DGGE fingerprinting implied that clustering was more related to the availability of organic carbon (the collapse of the P. gobosa bloom) than to the nature of the phytoplankton growth-controlling nutrient. Reduced irradiance delayed the development of the P. globosa population and subsequently changes in the bacterial community composition. Disintegration of P. globosa colonies resulted in the formation of transparent exopolymeric particles (TEP) and aggregates, more so under P-depletion than under N-deficient conditions. The colonial matrix transformed into big aggregates under P-depleted conditions but remained largely as ghost colonies under N-depleted conditions. In the mesocosm with initial nitrogen and phosphorus supplied in the Redfield ratio, features intermediate to conditions with either N- or P-depletion were observed. It was hypothesized that TEP affected microbial population dynamics directly through bacterial colonization and indirectly through scavenging of predators and viruses.     

The effects of oils and oil components on algae: A review

Plate structure analysis of the bloom-forming Peridinium of Lake Kinneret (previously identified as P. cinctum fa. westii) established its identity as P. gatunense. Peridinium cinctum was not observed. Material from different years and periods of the bloom, as well as specimens from cultures and P. gatunense from Lago Cristalino (Brazil), were studied using light and scanning electron microscopy. Peridinium gatunense from Lake Kinneret showed slight differences of the plate pattern as compared with specimens from other localities.     

Effect of nutrient loading and retention time on performance of high rate algal ponds

Small pilot ponds in a glasshouse at the Scottish Agricultural College (Auchincruive) were used to investigate the effects of changing C:N:P loading rate and retention time on pond performance as measured by nutrient removal and dry matter biomass. One experiment investigated ponds operated at two C:N:P ratios: low (9:7:1) and high (104:10:1) and two retention times (4 and 7 days θ. Increasing retention time from 4 to 7 days increased the concentration of total (dry matter) and algal (chlorophyll a) biomass and the degree of nitrification. It also increased removal of phosphorus, but had no effect on nitrogen or COD removal. Cyanobacteria predominated in ponds operated at both 4 and 7 days, and the density of cyanobacteria increased with increased retention time. Nitrogen removal was independent of C:N:P ratio; indeed the lower C:N:P ratio favoured increased nitrification. A high C:N:P ratio increased phosphorus and COD removal and increased the concentration of algal biomass (chlorophyll a), but had little effect on total biomass (dry matter). A second experiment varied COD loading rate (600, 350 and 100 kg COD ha-¹ d-¹) while maintaining a constant retention time (either 5 or 7 days θ). Species composition was independent of retention time. The longer retention time increased both total and algal biomass concentration and also percentage of nitrogen removed. Nitrification was independent of retention time. Increasing loading rate increased dry matter production and resulted in a predominance of cyanobacteria over Chlorophyceae. Increased loading rate was related to increase in nitrogen removal, however more complete nitrification occurred at low COD loading rates. Phosphorus removal in the pond with 5-day (θ) remained constant independent of loading rate, but in the pond with 7-day θ phosphorus removal increased with increased COD loading. COD removal was independent of both retention time and loading rate. This revised version was published online in August 2006 with corrections to the Cover Date.     

珠江口及毗邻海域营养盐对浮游植物生长的影响

基于2006年7月(夏季),10月(秋季)和2007年3月(春季)的现场调查数据,对珠江口及毗邻海域中的营养盐和叶绿素a等环境生态因子的时空分布特性进行了对比分析,研究了氮磷比与叶绿素a含量和种群多样性之间的联系,探讨了该海域营养盐对于浮游植物生长的影响。结果表明:(1)研究海域营养盐表现出较强的季节和空间差异性,总氮(TN)和总磷(TP)浓度均值春季(1.545 mg/L、0.056 mg/L)和夏季(1.570 mg/L、0.058 mg/L)均大于秋季(1.442 mg/L、0.034 mg/L),且春夏季浓度空间差异更明显。(2)调查期间海域营养盐含量超标现象突出,夏季尤为明显。无机氮(DIN)总体均值0.99 mg/L,超四类海水标准限值1倍,活性磷酸盐(PO4-P)总体均值0.021 mg/L,DIN∶PO4-P平均值为130;叶绿素a浓度与营养盐、p H、温度有较显著的相关性。(3)叶绿素a浓度较高的站位,具有较高的DIN∶PO4-P值,但浮游植物多样性指数偏低,优势种明显,主要为中肋骨条藻。氮磷比的改变会影响不同生长特性的浮游植物间的竞争和种群结构的改变;今后海洋污染治理中,在控制氮、磷污染时要注意氮磷比的改变可能造成的浮游生态影响。     

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.     

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 .     

Sulphate reduction in the hypolimnion and sediments of Lake Kinneret, Israel

1. Lake Kinneret is a warm (13–30°C) monomictic lake. Between January and June a heavy annual bloom of the dinoflagellate Peridinium gatunense dominates phytoplankton biomass (250 g m^?2). At the beginning of the summer, degradation and decomposition of the Peridinium biomass occurs, serving as a trigger for intense sulphate reduction in the hypolimnion and sediments. 2. The rates of sulphate reduction in the sediments varied seasonally from 12 to 1700 nmol SO₄.^?2 reduced cm^?3 day^?1 in December and July, respectively. The availability of organic matter and sulphate is high in June after the crash of the Peridinium bloom and the beginning of stratification and is lowest in December before overturn. 3. Sulphate concentrations in the hypolimnion range between 0.52 mM and 0.20 mM during mixing (January-April) and before overturn (December), respectively. The depletion in sulphate in the hypolimnion is stoichiometrically correlated to the increase in sulphide. The lake is not depleted of sulphate at any time, so the sulphate reduction process in Lake Kinneret is not limited by sulphate concentrations except in the sediments just before overturn.     

Lake Kinneret phytoplankton: Response to N and P enrichments in experiments and in nature

A series of water samples from Lake Kinneret was supplemented with 100 µM N (as NH₄ or as NO₃ and/or 10 µM orthophosphate-P. The yield of phytoplankton both as chlorophyll and in cell numbers of major species was determined after a two-week incubation. During these experiments, some of the algae present initially never multiplied (e.g. Peridinium and Peridiniopsis spp. Cryptomonas spp., Rhodomonas spp. and Crysochromulina parva); others e.g. Anomoeoneis exilis, Synedra sp., Chlamydomonas sp., Elakatothrix gelatinosa), undetected in the original sample, grew out during the incubation. Chlorophyte species (the majority of commonly observed forms in the lake) responded most readily to added nutrients.The results of these enrichment experiments were related to the long-term record of phytoplankton populations observed in the lake and suggest that through summer and fall, when ambient levels of both P and N are minimal, P was generally, but not always, the most limiting nutrient for algal growth. In the spring, after the decline of the Peridinium bloom, P appeared to be limiting the growth of Chlorophyta. Although most algal species grew equally well on NH₄ or NO₃, some species appeared to respond preferentially either to the former (Coelastrum, Chodatella) or to the latter (Chroococcus, Anomoeoneis) source of N.     

RESPONSE OF A LENTIC PERIPHYTON COMMUNITY TO NUTRIENT ENRICHMENT AT LOW N:P RATIOS1

We examined the effect of nitrogen:phosphorus (N:P) ratios and nutrient concentrations on periphyton when nutrients (N and P) are provided in excess. A gradient of seven N:P ratios ranging from 7.5:1 to 1:7.5 and each at three absolute concentrations, was established using nutrient‐releasing substrata placed in a meso‐oligotrophic lake. Differences in total algal biovolume among nutrient ratios were significant (analysis of covariance [ANCOVA]) when P concentration was entered as the co‐variate. In addition, total algal biovolume was significantly correlated with N concentration but not P. To further evaluate the relationship between nutrient ratios and biovolume, we analyzed (using four 1‐way analysis of variances [ANOVAs]) four subsets of data defined as a series of treatments where one nutrient concentration remained relatively constant as the other changed creating different N:P ratios. Ratios of data subsets ranged from 1:1 to 7.5:1 and 1:1 to 1:7.5 with low and high concentrations of both series. Only diatom biovolume varied with ratio but these differences are most likely related to increased green algal abundance. Species richness and diversity differed among N:P ratios (ANCOVA) when P concentration was used as the co‐variate. Stigeoclonium tenue (Ag.) Gomont, which generally accounted for the increase in green algal abundance, varied with nutrient ratio (ANCOVA) when P was the co‐variate. Based on the ANCOVAs, correlations, and one‐way ANOVAs, periphyton in this system appears to be affected by N concentration but not by N:P treatment ratios under nutrient‐rich conditions. When compared with previous studies, these data also suggest that the response of periphyton to in situ treatments constructed with nutrient‐releasing substrata vary between years.     

Uncoupling of chlorophyll and nutrients in lakes – possible reasons,expected consequences

Total phosphorus and total nitrogen explained a low percentage of summer chlorophyll variability in epilimnia of the Great Masurian Lakes. Division of the whole data set into two subgroups of lakes improved approximation of the chlorophyll nutrient relationship but revealed also functional differences between the lakes distinguished in that way. Chlorophyll in eutrophic lakes correlated well with nitrogen and phosphorus, that in mesotrophic lakes (those with summer chlorophyll <=22 mg m^–3 as calculated in the model) was related to none of the nutrients. Higher summer chlorophyll content in epilimnetic waters was accompanied by higher chl:PP and chl:PN ratios. Algal adaptation to poor light conditions in eutrophic lakes is postulated as a possible reason for that difference.Chlorophyll – nutrient relationships varied with the trophic status of lakes. Epilimnetic chlorophyll strictly followed phosphorus changes in eutrophic lakes but did not do so in mesotrophic ones. Detailed comparison of selected meso- and eutrophic lakes showed marked differences in the seasonal changes of chlorophyll and nutrient concentrations and in sedimentation rates, especially in spring. Nutrient limitation rather than zooplankton grazing is suggested as a possible mechanism of controlling algal abundance and the sequence of spring events in a eutrophic lake. It is hypothesised that phosphorus turnover in eutrophic lakes is dominated by seasonal vertical fluxes, while in mesotrophic lakes it is more conservative with consumption and regeneration restricted mostly to metalimnion. Possible consequences of such conclusion are discussed in the paper.     

Effects of zooplankton grazing and nutrients on the bloom-forming, N2-fixing cyanobacterium Aphanizomenon in Lake Kinneret

A bloom of the filamentous, N₂-fixing cyanobacterium Aphanizomenonovalisporum Forti occurred for the first time in Lake Kinneretduring late summer and fall 1994. During subsequent years (1995–1999),Aphanizomenon also appeared in late summer and fall, but didnot bloom. In outdoor microcosm experiments, we examined zooplanktongrazing on Lake Kinneret phytoplankton, with and without Aphanizomenonpresent, and the effects of N, P and N:P ratios on phytoplanktongrowth. In one-day feeding experiments, clearance and grazingrates of the ambient Lake Kinneret zooplankton assemblage feedingin lake water dominated by Aphanizomenon were 10-fold lowerthan in water without Aphanizomenon. We suspect that the lowgrazing rates were due to interference caused by the presenceof Aphanizomenon. In 9-day nutrient addition experiments, significantenhancement effects on phytoplankton were detected with additionsof either P or N; a high N:P was better for phytoplankton growththan a low N:P. After 7 days, bottles receiving low P and noN additions were dominated by Oscillatoria sp. and Closteriumacutum; few Aphanizomenon were present. In contrast, bottlesreceiving high P and N additions had large increases of Aphanizomenon,as well as Oscillatoria and Closterium. There was a tendencyfor more green algae and diatoms with increasing N additions.These results provide evidence that (i) non-grazeability ofAphanizomenon enabled it to gain a competitive advantage overgrazeable phytoplankton, and (ii) that nutrient limitation,but not grazing, was probably important in the eventual declineof the Aphanizomenon bloom.     

Changes to the phytoplankton assemblage of Lake Kinneret after decades of a predictable, repetitive pattern

1. Phytoplankton abundance and species composition in Lake Kinneret, Israel, have been monitored at weekly or fortnightly intervals since 1969. This paper summarises the resulting 34‐year phytoplankton record with a focus on the last 13 years of new data, and reassesses an earlier conclusion that the lake phytoplankton shows remarkable stability despite a wide range of external pressures. 2. The Kinneret phytoplankton record can be split into two major periods. The first, from 1969 till 1993, was a period of distinct stability expressed by a typical annual pattern revolving around a spring bloom of the dinoflagellate Peridinium gatunense that repeated each year. The second period, starting around 1994 and ongoing, is characterised by the loss of the previously predictable annual pattern, with both ‘bloom years’ and ‘no‐bloom years’. 3. In the second period, deviations from the previous annual pattern include: the absence of the prevailing spring P. gatunense blooms in some years and increased variability in the magnitude of the bloom in others; intensification of winter Aulacoseira granulata blooms; higher summer phytoplankton biomass with replacement of mostly nanoplanktonic, palatable forms by less palatable forms; new appearance and establishment of toxin‐producing, nitrogen fixing cyanobacteria in summer; increase in the absolute biomass and percentage contribution of cyanobacteria to total biomass; and fungal epidemics attacking P. gatunense. 4. The 34‐year record serves to validate Schindler's (1987) assessment that phytoplankton species composition will respond to increased anthropogenic stress before bulk ecosystem parameters.     

施肥与刈割对内蒙古草地生态化学计量特征的影响

为明确施肥和刈割对蒙古半干旱草地四种植物化学计量特征的影响,该文于2014年8月以内蒙古半干旱草地的四种植物冷蒿(Artimesia frigida)、菊叶委陵菜(Potentilla tanacetifolia)、羊草(Aneurotepidimu chinense)、扁蓿豆(Melissitus ruthenica)为研究对象,利用方差分析及多重比较分析对照、施氮肥、割草、割草和施氮肥处理下的植物碳、氮、磷含量的差异以及不同处理下碳氮比和氮磷比的变化。结果表明:施氮肥与割草均未影响植物碳含量。施氮肥显著提高了植物氮含量(P<0.05),进而降低了碳氮比(P<0.05),提高了氮磷比(P<0.05)。割草处理后扁蓿豆的氮含量显著提高了0.18%(P<0.05),羊草的氮含量显著降低了0.13%(P<0.05),冷蒿与菊叶委陵菜的氮含量无显著变化。割草显著提高了冷蒿和菊叶委陵菜的磷含量(P<0.05),但是未影响羊草和扁蓿豆的磷含量。该研究证实了氮添加或割草会影响部分草地物种的化学计量特征,但是氮添加或割草对植物化学计量特征的影响存在种间差异。     

The C:N:P ratio of phytoplankton determines the relative amounts of dissolved inorganic nitrogen and phosphorus released during aerobic decomposition

Three phytoplankton assemblages, with different C:N:P ratios of 314:55:1, 103:17:1, and 57:5.5:1 (by weight), were prepared by growing lake phytoplankton in artificial media with different N:P supply ratios and then decomposed under aerobic conditions for three months.There was no net release of dissolved inorganic phosphorus (DIP) from the phytoplankton assemblage with the highest C:N:P ratio, in contrast with an abundant liberation of DIP from that with the lowest C:N:P ratio. On the other hand, there was an abundant release of dissolved inorganic nitrogen (DIN) from the phytoplankton assemblage with the highest C:N:P ratio, in contrast with little or virtually no release of DIN from those with lower C:N:P ratios. Thus, it was concluded that the C:N:P ratio of phytoplankton is an important parameter to determine the relative amounts of DIP and DIN released, when they are decomposed under aerobic conditions.Contribution from Otsu Hydrobiological Station, Kyoto University, No. 316 (Foreign Language Series).Contribution from Otsu Hydrobiological Station, Kyoto University, No. 316 (Foreign Language Series).     

A mesocosm study of Phaeocystis globosa population dynamics: I. Regulatory role of viruses in bloom control

The regulatory role of viruses on population dynamics of the prymnesiophyte Phaeocystis globosa was studied during a mesocosm experiment in relation to growth and loss by microzooplankton grazing and cell lysis. The mesocosms were conducted under varying light conditions (20 and 150 μmol photons m⁻² s⁻¹) and nutrient regime (inorganic nitrogen to phosphorus ratios of 4, 16 and 44). Overall, viruses infecting P. globosa (PgV) were found to be an important cause of cell lysis (30–100% of total lysis) and a significant loss factor (7–67% of total loss). We demonstrate that the morphology of P. globosa cells (solitary versus colonial) differently regulated viral control of P. globosa bloom formation. Reduced irradiance (20 μmol photons m⁻² s⁻¹) was provided for 11 days to select for the solitary cell morphotype. Viruses were able to restrict P. globosa bloom formation even after irradiance became saturating again (150 μmol photons m⁻² s⁻¹). Saturating light conditions from the start of the experiment allowed colony formation and because the colony-morphotype acted as a mechanism reducing viral infection bloom formation succeeded. Nutrient depletion, however, affected specifically the colonies that disintegrated while releasing single cells. Virus infection of these solitary cells resulted in the termination of the bloom. The nature of phytoplankton growth-limiting nutrient (nitrate and/or orthophosphate) did not seem to noticeably affect the level of viral control.     

Inter- and intra-annual variability in the phytoplankton community of a high mountain lake: the influence of external (atmospheric) and internal (recycled) sources of phosphorus

1. The inter‐ and intra‐annual changes in the biomass, elemental (carbon (C), nitrogen (N) and phosphorus (P)) and taxonomical composition of the phytoplankton in a high mountain lake in Spain were studied during 3 years with different physical (fluctuating hydrological regime) and chemical conditions. The importance of internal and external sources of P to the phytoplankton was estimated as the amount of P supplied via zooplankton recycling (internal) or through ice‐melting and atmospheric deposition (external). 2. Inter‐annual differences in phytoplankton biomass were associated with temperature and total dissolved phosphorus. In 1995, phytoplankton biomass was positively correlated with total dissolved phosphorus. In contrast, the negative relationship between zooplankton and seston biomass (direct predatory effects) and the positive relationship between zooplankton P excretion and phytoplankton biomass in 1997 (indirect P‐recycling effects), reinforces the primary role of zooplankton in regulating the total biomass of phytoplankton but, at the same time, encouraging its growth via P‐recycling. 3. Year‐to‐year variations in seston C : P and N : P ratios exceeded intra‐annual variations. The C : P and N : P ratios were high in 1995, indicating strong P limitation. In contrast, in 1996 and 1997, these ratios were low during ice‐out (C : P < 100 and N : P < 10) and increased markedly as the season progressed. Atmospheric P load to the lake was responsible for the decline in C : P and N : P ratios. 4. Intra‐annual variations in zooplankton stoichiometry were more pronounced than the overall differences between 1995 and 1996. Thus, the zooplankton N : P ratio ranged from 6.9 to 40.1 (mean 21.4) in 1995, and from 10.4 to 42.2 (mean 24.9) in 1996. The zooplankton N : P ratio tended to be low after ice‐out, when the zooplankton community was dominated by copepod nauplii, and high towards mid‐ and late‐season, when these were replaced by copepodites and adults. 5. In 1995, the minimum demands for P of phytoplankton were satisfied by ice‐melting, atmospheric loading and zooplankton recycling over 100%. In order of importance, atmospheric inputs (> 1000%), zooplankton recycling (9–542%), and ice‐melting processes (0.37–5.16%) satisfied the minimum demand for P of phytoplankton during 1996 and 1997. Although the effect of external forces was rather sporadic and unpredictable in comparison with biologically driven recycle processes, both may affect phytoplankton structure and elemental composition. 6. We identified three conceptual models representing the seasonal phosphorus flux among the major compartments of the pelagic zone. While ice‐melting processes dominated the nutrient flow at the thaw, biologically driven processes such as zooplankton recycling became relevant as the season and zooplankton ontogeny progressed. The stochastic nature of P inputs associated with atmospheric events can promote rapid transitional changes between a community limited by internal recycling and one regulated by external load. 7. The elemental composition of the zooplankton explains changes in phytoplankton taxonomic and elemental composition. The elemental negative balance (seston N : P < zooplankton N : P, low N : P recycled) during the thaw, would promote a community dominated by species with a high demand for P (Cryptophyceae). The shift to an elemental positive balance (seston N : P > zooplankton N : P, high N : P recycled) in mid‐season would skew the N : P ratio of the recycled nutrients, favouring dominance by chrysophytes. The return to negative balance, as a consequence of the ontogenetic increase in zooplankton N : P ratio and the external P inputs towards the end of the ice‐free season, could alleviate the limitation of P and account for the appearance of other phytoplankton classes (Chlorophyceae or Dinophyceae).     

Phosphorus inputs unmask negative effects of ultraviolet radiation on algae in a high mountain lake

Ultraviolet solar radiation (UVR) and atmospheric nutrient loads to pristine ecosystems are global climate change phenomena that simultaneously affect aquatic organisms in ways not easily predicted by single factor studies. Plankton in a high mountain lake was exposed in situ to increasing phosphorus (P) concentrations (mimicking atmospheric pulses) in absence or presence of UVR in order to identify their interactive effect on functional [primary production, organic carbon (C) release (EOC), and percentage of C released (%EOC)], growth rate, structural–physiological (algal biomass, sestonic C, P content, chlorophyll a (Chl a), and Chl a : C ratio, P cell quota, cell‐specific Chl a), and stoichiometric (autotroph C : P ratio) traits. The availability of P after the pulse determined the intensity of responses by primary producers to UVR stress. All structural–physiological and functional variables significantly increased by up to two orders of magnitude in response to P enrichment. UV radiation, over a long‐term scale, exerted significant deleterious effects on most structural–physiological variables when inorganic P was added at high levels (≥30 μg P L^?1). The subsequent unexpected negative synergistic UVR × P effect on algal development did not support our initial hypothesis that P input might buffer the harmful UVR effect. UVR exerted a weak negative effect on primary production but strongly enhanced the absolute and percentage excretion of C (up to 60%), mechanism responsible of a significant reduction in autotroph C : P ratios. We propose that low sestonic C : P ratios are the outcome of an adaptive strategy of algae in environments with high UVR exposure and extreme nutrient limitation and have important implications for C flux through grazing vs. microbial food webs in oligotrophic systems.