Burial rates and stoichiometry of sedimentary carbon,nitrogen and phosphorus in Midwestern US reservoirs

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Patterns of seasonal phytoplankton distribution in prairie saline lakes of the northern Great Plains (U.S.A.)

Seasonal changes in freshwater phytoplankton communities have been extensively studied, but key drivers of phytoplankton in saline lakes are currently not well understood. Comparative lake studies of 19 prairie saline lakes in the northern Great Plains (USA) were conducted in spring and summer of 2004, with data gathered for a suite of limnological parameters. Nutrient enrichment assays for natural phytoplankton assemblages were also performed in spring and summer of 2006. Canonical correspondence analysis of 2004 data showed salinity (logCl), nitrogen, and phosphorus (N:P ratios) to be the main drivers of phytoplankton distribution in the spring, and phosphorus (C:P ratios), iron (logTFe), and nitrogen (logTN) as important factors in the summer. Despite major differences in nutrient limitation patterns (P-limitation in freshwater systems, N-limitation in saline systems), seasonal patterns of phytoplankton phyla changes in these saline lakes were similar to those of freshwater systems. Dominance shifted from diatoms in the spring to cyanobacteria in the summer. Nutrient enrichment assays (control, +Fe, +N, +P, +N+P) in 2006 indicated that nutrient limitation is generally more consistent within lakes than for individual taxa across systems, with widespread nitrogen and secondary phosphorus limitation. Understanding phytoplankton community structure provides insight into the overall ecology of saline lakes, and will assist in the future conservation and management of these valuable and climatically-sensitive systems.     

Chemical composition of modern and pre-acidification sediments in the Tatra Mountain lakes

Concentrations of major nutrients (C, N, P) and acid soluble metals (Ca, Mg, K, Al, Fe, Mn, Pb, and Zn) were determined in modern (0–1 cm) and pre-acidification (5–10 cm) sediment layers collected from 37 alpine and 3 forest lakes in the Tatra Mountains (Slovakia, Poland) in 1996–1998. Sediment composition reflected catchment characteristics and productivity of lakes. In the sediments of alpine lakes, C and N concentrations decreased and Mg increased with a decreasing proportion of vegetation and soil in the catchment. Decreasing Ca:Mg ratios in sediments along the vegetation gradient was inverse to that in water, and could be associated with different ratios of cations in water leachate from catchments and in solids which enter the lake due to soil erosion. Phosphorus concentrations increased with the proportion of moraine areas, with till soils rich in P. Concentrations of C, N, P, and Ca in sediments positively correlated to their concentrations in water. Sediment concentrations of Al and Al:Ca ratios increased with decreasing sediment and water pH. A negative correlation between water pH and concentrations of organic C in water and sediments indicated the important impact of organic acids on the acid status of the lakes exposed to higher terrestrial export of organic matter. Compared to the pre-acidification period, the modern sediments had significantly higher Fe, Mn, Zn, Pb, and K, but lower Mg concentrations. The Zn and Pb enrichment was more evident in oligotrophic alpine lakes than in more productive forest lakes and was independent of lake water or sediment pH. Fe and Mn concentrations in the modern sediments were higher than in ambient soils and bedrock, while those in pre-acidification sediments were similar to contemporary soils and to the rock layer. The enrichment of the modern sediments with Fe and Mn thus probably resulted from both their redox recycling and ecosystem acidification.     

Phytoplankton growth and stoichiometric responses to warming,nutrient addition and grazing depend on lake productivity and cell size

Global change involves shifts in multiple environmental factors that act in concert to shape ecological systems in ways that depend on local biotic and abiotic conditions. Little is known about the effects of combined global change stressors on phytoplankton communities, and particularly how these are mediated by distinct community properties such as productivity, grazing pressure and size distribution. Here, we tested for the effects of warming and eutrophication on phytoplankton net growth rate and C:N:P stoichiometry in two phytoplankton cell size fractions (<30 µm and >30 µm) in the presence and absence of grazing in microcosm experiments. Because effects may also depend on lake productivity, we used phytoplankton communities from three Dutch lakes spanning a trophic gradient. We measured the response of each community to multifactorial combinations of temperature, nutrient, and grazing treatments and found that nutrients elevated net growth rates and reduced carbon:nutrient ratios of all three phytoplankton communities. Warming effects on growth and stoichiometry depended on nutrient supply and lake productivity, with enhanced growth in the most productive community dominated by cyanobacteria, and strongest stoichiometric responses in the most oligotrophic community at ambient nutrient levels. Grazing effects were also most evident in the most oligotrophic community, with reduced net growth rates and phytoplankton C:P stoichiometry that suggests consumer‐driven nutrient recycling. Our experiments indicate that stoichiometric responses to warming and interactions with nutrient addition and grazing are not universal but depend on lake productivity and cell size distribution.     

Seasonal Patterns of Nitrogen and Phosphorus Limitation in Four German Lakes and the Predictability of Limitation Status from Ambient Nutrient Concentrations

To identify the seasonal pattern of nitrogen (N) and phosphorus (P) limitation of phytoplankton in four different lakes, biweekly experiments were conducted from the end of March to September 2011. Lake water samples were enriched with N, P or both nutrients and incubated under two different light intensities. Chlorophyll a fluorescence (Chla) was measured and a model selection procedure was used to assign bioassay outcomes to different limitation categories. N and P were both limiting at some point. For the shallow lakes there was a trend from P limitation in spring to N or light limitation later in the year, while the deep lake remained predominantly P limited. To determine the ability of in-lake N:P ratios to predict the relative strength of N vs. P limitation, three separate regression models were fit with the log-transformed ratio of Chla of the P and N treatments (Response ratio = RR) as the response variable and those of ambient total phosphorus:total nitrogen (TN:TP), dissolved inorganic nitrogen:soluble reactive phosphorus (DIN:SRP), TN:SRP and DIN:TP mass ratios as predictors. All four N:P ratios had significant positive relationships with RR, such that high N:P ratios were associated with P limitation and low N:P ratios with N limitation. The TN:TP and DIN:TP ratios performed better than the DIN:SRP and TN:SRP in terms of misclassification rate and the DIN:TP ratio had the highest R² value. Nitrogen limitation was predictable, frequent and persistent, suggesting that nitrogen reduction could play a role in water quality management. However, there is still uncertainty about the efficacy of N restriction to control populations of N₂ fixing cyanobacteria.     

Interactive effects of light and nutrients on phytoplankton stoichiometry

The stoichiometric composition of autotrophs can vary greatly in response to variation in light and nutrient availability, and can mediate ecological processes such as C sequestration, growth of herbivores, and nutrient cycling. We investigated light and nutrient effects on phytoplankton stoichiometry, employing five experiments on intact phytoplankton assemblages from three lakes varying in productivity and species composition. Each experiment employed two nutrient and eight irradiance levels in a fully factorial design. Light and nutrients interactively affected phytoplankton stoichiometry. Thus, phytoplankton C:N, C:P, and N:P ratios increased with irradiance, and slopes of the stoichiometric ratio versus irradiance relationships were steeper with ambient nutrients than with nutrients added. Our results support the light–nutrient hypothesis, which predicts that phytoplankton C:nutrient ratios are functions of the ratio of available light and nutrients; however, we observed considerable variation among lakes in the expression of this relationship. Phytoplankton species diversity was positively correlated with the slopes of the C:N and C:P versus irradiance relationships, suggesting that diverse assemblages may exhibit greater flexibility in the response of phytoplankton nutrient stoichiometry to light and nutrients. The interactive nature of light and nutrient effects may render it difficult to generate predictive models of stoichiometric responses to these two factors. Our results point to the need for future studies that examine stoichiometric responses across a wide range of phytoplankton communities.     

Elemental ratios and the uptake and release of nutrients by phytoplankton and bacteria in three lakes of the Canadian shield

The dynamics of carbon (C), nitrogen (N), and phosphorus (P), elemental ratios, and dark uptake/release of N and P in bacterial and phytoplankton size fractions were studied during summer 1992 in three lakes of contrasting food web structure and trophic status (L240, L110, L227). We wished to determine if phytoplankton and bacteria differed in their elemental characteristics and to evaluate whether the functional role of bacteria in nutrient cycling (i.e., as sink or source) depended on bacterial elemental characteristics. Bacterial contributions to total suspended particulate material and to fluxes of nutrients in the dark were substantial and varied for different elements. This indicated that some techniques for assaying phytoplankton physiological condition are compromised by bacterial contributions. C/N ratios were generally less variable than C/P and N/P ratios. Both elemental ratios and biomass-normalized N and P flux indicated that phytoplankton growth in each lake was predominantly P-limited, although in L227 these data reflect the dominance of N-fixing cyanobacteria, and N was likely limiting early in the sampling season. In L227, phytoplankton N/P ratio and biomass-normalized N flux were negatively correlated, indicating that flux data were likely a reasonable measure of the N status of the phytoplankton. However, for L227 phytoplankton, P-flux per unit biomass was a hyperbolic function of N/P, suggesting that the dominant L227 cyanobacteria have a limited uptake and storage capacity and that P-flux per unit biomass may not be a good gauge of the P-limitation status of phytoplankton in this situation. Examination of N-flux data in the bacterial size fraction relative to the N/P ratio of the bacteria revealed a threshold N/P ratio (22:1 N/P, by atoms), below which, bacteria took up and sequestered added N, and above which, N was released. Thus, the functional role of bacteria in N cycling in these ecosystems depended on their N/P stoichiometry.     

Nitrogen deposition and warming – effects on phytoplankton nutrient limitation in subarctic lakes

The aim of this study was to predict the combined effects of enhanced nitrogen (N) deposition and warming on phytoplankton development in high latitude and mountain lakes. Consequently, we assessed, in a series of enclosure experiments, how lake water nutrient stoichiometry and phytoplankton nutrient limitation varied over the growing season in 11 lakes situated along an altitudinal/climate gradient with low N‐deposition (<1 kg N ha^?1 yr^?1) in northern subarctic Sweden. Short‐term bioassay experiments with N‐ and P‐additions revealed that phytoplankton in high‐alpine lakes were more prone to P‐limitation, and with decreasing altitude became increasingly N‐ and NP‐colimited. Nutrient limitation was additionally most obvious in midsummer. There was also a strong positive correlation between phytoplankton growth and water temperature in the bioassays. Although excess nutrients were available in spring and autumn, on these occasions growth was likely constrained by low water temperatures. These results imply that enhanced N‐deposition over the Swedish mountain areas will, with the exception of high‐alpine lakes, enhance biomass and drive phytoplankton from N‐ to P‐limitation. However, if not accompanied by warming, N‐input from deposition will stimulate limited phytoplankton growth due to low water temperatures during large parts of the growing season. Direct effects of warming, allowing increased metabolic rates and an extension of the growing season, seem equally crucial to synergistically enhance phytoplankton development in these lakes.     

Phytoplankton nutrient limitation in Colorado mountain lakes

SUMMARY. 1. Limiting nutrients for phytoplankton were studied experimentally in eight mountain lakes of central Colorado between May and November of 1984.
2. Five categories of phytoplankton limitation were identified: no limitation, N limitation, P limitation, concurrent limitation (stimulation only by simultaneous additions of N and P), and reciprocal limitation (stimulation by addition of either N or P). The phytoplankton communities of three lakes were primarily N-limited, one was primarily phosphorus-limited, and four showed primarily combined limitation (concurrent or reciprocal). Switching between categories of limitation was also observed within lakes. Nitrogen was the most frequently limiting nutrient; N, either alone or in combination with P, accounted for 79% of all observed instances of limitation.
3. Nine indices were tested for effectiveness in predicting phytoplankton limitation by N and P. The best indices for discriminating all limitations were ratios of dissolved inorganic N: total P (84% accuracy) and dissolved inorganic N:total dissolved P (80% accuracy). The effectiveness of these indices may be explained by the degree to which they represent N and P fractions actually available to the phytoplankton.     

Growth rate of alpine phytoplankton assemblages from contrasting watersheds and N‐deposition regimes exposed to nitrogen and phosphorus enrichments

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Atmospheric nitrogen-deposition may intensify phosphorus limitation of shallow epilithic periphyton in unproductive lakes

1. We tested whether increasing atmospheric nitrogen (N) deposition along a north–south gradient intensifies epilithic phosphorus (P) limitation in oligotrophic Swedish lakes from the north to the south. We examined the epilithic community at a shallow depth from seven northern and six southern Swedish lakes, and also compared the results with a lake located geographically between the two groups. We determined lake nutrient state, epilithic nutrient ratios and epilithic algal composition, as well as grazer N : P ratios, grazer-epilithon N : P imbalance, and N : P cycling ratios.
2. Epilithic communities appear to be generally more N-limited in the northern lakes and more P-limited in the southern lakes. Lake water total N (Tot-N) and epilithic N : P ratios were lower in northern than in southern lakes and the proportion of N₂-fixing cyanobacteria was higher in northern than in southern lakes.
3. Gastropod grazers had lower N : P imbalances and cycled less N relative to P in northern than in southern lakes.
4. Atmospheric N-deposition showed a strong positive correlation with lake water Tot-N and a much weaker positive correlation with epilithon N : P ratios. Atmospheric N-deposition also correlated negatively with the proportion of N₂-fixing cyanobacteria.
5. There are indications that increased atmospheric N-deposition towards the south might intensify P-limitation of epilithic algae and invertebrate grazers, although more studies are needed to show the strength and generality of our findings.     

Sediment microbial enzyme activity as an indicator of nutrient limitation in Great Lakes coastal wetlands

1. We compared the extracellular enzyme activity (EEA) of sediment microbial assemblages with sediment and water chemistry, gradients in agricultural nutrient loading (derived from principal component analyses), atmospheric deposition and hydrological turnover time in coastal wetlands of the Laurentian Great Lakes. 2. There were distinct increases in nutrient concentrations in the water and in atmospheric N deposition along the gradient from Lake Superior to Lake Ontario, but few differences between lakes in sediment carbon (C), nitrogen (N) or phosphorus (P). Wetland water and sediment chemistry were correlated with the agricultural stress gradient, hydrological turnover time and atmospheric deposition. 3. The N : P ratio of wetland waters and sediments indicated that these coastal wetlands were N‐limited. Nutrient stoichiometry was correlated with the agricultural stress gradient, hydrological turnover time and atmospheric deposition. 4. Extracellular enzyme activity was correlated with wetland sediment and water chemistry and stoichiometry, atmospheric N deposition, the agricultural stress gradient and the hydrological turnover time. The ratios of glycosidases to peptidases and phosphatases yielded estimates of nutrient limitation that agreed with those based solely on nutrient chemistry. 5. This study, the first to link microbial enzyme activities to regional‐scale anthropogenic stressors, suggests that quantities and ratios of microbial enzymes are directly related to the concentrations and ratios of limiting nutrients, and may be sensitive indicators of nutrient dynamics in wetland ecosystems, but further work is needed to elucidate these relationships.     

Nitrogen mass balances and denitrification rates in central Ontario Lakes

Nitrogen mass balances for seven unproductive lakes and 20 forested catchments in central Ontario were measured between 1977 and 1989. Average annual lake denitrification rates calculated with the N/P ratio method were strongly correlated with summer anoxic factor (extent of surficial sediment anoxia) whereas denitrification rates calculated with a²¹⁰Pb sediment N accumulation method were poorly correlated with the anoxic factor suggesting that the N/P method is superior. Substantial denitrification occurred in all lakes — an average of 36% of TN inputs or 75% of the net gain. On a regional area-weighted basis, 67% of bulk atmospheric TN deposition was stored or denitrified terrestrially, 12% was denitrified in lakes, 4% was stored in lake sediments, and 17% was exported from lakes. N/P ratios were generally less in streams than in precipitation suggesting preferential N retention in catchments, whereas the N/P ratios in lake outputs were slightly higher than lake input ratios, suggesting preferential P retention in lakes. This is consistent with the notion that P-limited lakes can exist adjacent to N-limited forests.     

Increased nutrient loading and rapid changes in phytoplankton expected with climate change in stratified South European lakes: sensitivity of lakes with different trophic state and catchment properties

We hypothesised that increasing winter affluence and summer temperatures, anticipated in southern Europe with climate change, will deteriorate the ecological status of lakes, especially in those with shorter retention time. We tested these hypotheses analysing weekly phytoplankton and chemistry data collected over 2 years of contrasting weather from two adjacent stratified lakes in North Italy, differing from each other by trophic state and water retention time. Dissolved oxygen concentrations were higher in colder hypolimnia of both lakes in the second year following the cold winter, despite the second summer was warmer and the lakes more strongly stratified. Higher loading during the rainy winter and spring increased nutrient (N, P, Si) concentrations, and a phytoplankton based trophic state index, whilst the N/P ratio decreased in both lakes. The weakened Si limitation in the second year enabled an increase of diatom biovolumes in spring in both lakes. Chlorophyll a concentration increased in the oligo-mesotrophic lake, but dropped markedly in the eutrophic lake where the series of commonly occurring cyanobacteria blooms was interrupted. The projected increase of winter precipitation in southern Europe is likely to increase the nutrient loadings to lakes and contribute to their eutrophication. The impact is proportional to the runoff/in-lake concentration ratio of nutrients rather than to the retention time, and is more pronounced in lakes with lower trophy.     

A worldwide view of organic carbon export from catchments

Growing interest in the effects of global change on the metabolism, stoichiometry and cycling of carbon in aquatic ecosystems has motivated research on the export of organic carbon (OCE) from catchments. In this article, quantitative and functional features of the annual export rates of total, particulate and dissolved organic carbon (TOC, POC and DOC) were reviewed, and the stoichiometry of export (OC:N, OC:P and N:P) from 550 catchments worldwide was reported. TOC export ranged 2.1–92,474?kg?C?km^?2?year^?1, POC export ranged 0.4–73,979?kg?C?km^?2?year^?1 and DOC export ranged 1.2–56,946?kg?C?km^?2?year^?1. Exports of TOC and DOC were strongly linked, but POC export was unrelated to DOC. The DOC fraction comprised on average 73?±?21% of TOC export. The export rates of organic carbon were poorly related to those of total nitrogen and total phosphorus. Discrete and continuous environmental variables failed to predict TOC export, but DOC export was influenced by discharge and catchment area worldwide. Models of OCE in different catchment types were controlled by different environmental variables; hydrological variables were generally better predictors of OCE than anthropogenic and soil variables. Elemental ratios of carbon export in most catchments were above the Redfield ratio, suggesting that phosphorus may become the limiting nutrient for downstream plant growth. These ratios were marginally related to environmental data. More detailed hydrological data, consideration of in-stream processes and the use of quasi-empirical dynamical models are advocated to improve our knowledge of OCE rates and those of other nutrients.     

Detritivory and the stoichiometry of nutrient cycling by a dominant fish species in lakes of varying productivity

Little is known about the stoichiometry of nutrient cycling by detritivores. Therefore, we explored stoichiometric relationships in an omnivorous/detritivorous fish (gizzard shad, Dorosoma cepedianum) in three lakes that differed in productivity. Gizzard shad can feed on plankton and sediment detritus, but in all three lakes adult gizzard shad derived >98% of carbon (C) and phosphorus (P), and >90% of nitrogen (N) from sediment detritus, and the remainder from zooplankton.
Gizzard shad selectively consumed detritus with higher C, N and P concentrations than ambient lake sediments. Selective detritivory (i.e. the nutrient content of consumed detritus divided by the nutrient content of ambient detritus) was most pronounced in the lake with the lowest detrital nutrient concentrations. N and P cycling rates per fish were also consistently higher in this lake, in agreement with the prediction of stoichiometry theory that excretion rates should increase with food nutrient content. Among-lake differences in nutrient cycling rates were unrelated to inter-lake variation in fish body nutrient contents, which was minimal. The N:P ratio excreted was near Redfield (∼14:1) in all three lakes.
Stoichiometric analyses showed that the C:N and C:P ratios of sediment detritus were much higher (∼2.8×) than ratios of gizzard shad bodies, revealing substantial N and P imbalances between consumers and their food source. Gizzard shad alleviate N imbalance by selectively feeding on high N detritus (low C:N, high N:P), and apparently alleviate P imbalance by excreting nutrients at a higher N:P than that of their food or their bodies. Thus, this detritivore apparently regulates nutrient acquisition and allocation via both pre-absorption processes (selective feeding) and post-absorptive processes (differential N and P excretion).     

Iron Constraints on Planktonic Primary Production in Oligotrophic Lakes

Phototrophic primary production is a fundamental ecosystem process, and it is ultimately constrained by access to limiting nutrients. Whereas most research on nutrient limitation of lacustrine phytoplankton has focused on phosphorus (P) and nitrogen (N) limitation, there is growing evidence that iron (Fe) limitation may be more common than previously acknowledged. Here we show that P was the nutrient that stimulated phytoplankton primary production most strongly in seven out of nine bioassay experiments with natural lake water from oligotrophic clearwater lakes. However, Fe put constraints on phytoplankton production in eight lakes. In one of these lakes, Fe was the nutrient that stimulated primary production most, and concurrent P and Fe limitation was observed in seven lakes. The effect of Fe addition increased with decreasing lake water concentrations of total phosphorus and dissolved organic matter. Possible mechanisms are low import rates and low bioavailability of Fe in the absence of organic chelators. The experimental results were used to predict the relative strength of Fe, N, and P limitation in 659 oligotrophic clearwater lakes (with total phosphorus ≤ 0.2 μM P and total organic carbon < 6 mg C l⁻¹) from a national lake survey. Fe was predicted to have a positive effect in 88% of these lakes, and to be the nutrient with the strongest effect in 30% of the lakes. In conclusion, Fe, along with P and N, is an important factor constraining primary production in oligotrophic clearwater lakes, which is a common lake-type throughout the northern biomes. This paper is dedicated to the memory of Prof. Peter Blomqvist (deceased 2004).     

Impact of DOC trends resulting from changing climatic extremes and atmospheric deposition chemistry on periphyton community of a freshwater tropical lake of India

Recent measurements have demonstrated unprecedented increase in atmospheric deposition of nutrients in many parts of India. To determine whether atmospheric nutrient inputs would increase phytoplankton growth and catchment dissolved organic carbon (DOC) flushing to constrain benthic algae, we analyzed NO₃ ^? and PO ₄ ^?3 in atmospheric deposits; nutrients and DOC in runoff and lake water and standing crop biomass of phytoplankton and periphyton at Jaisamand Lake of Rajasthan, India. Atmospheric deposition of NO₃ ^? (7.18–29.95 kg ha^?1 year^?1) and PO ₄ ^?3 (0.56–2.15 kg ha^?1 year^?1) showed a consistently rising trend across the year. Microbial biomass and activity in catchment increased in response to atmospheric deposition. Lake DOC and nutrients showed strong coherence with their terrestrial and atmospheric fluxes. Phytoplankton development showed significant linearity with atmospheric input of nutrients. Air-driven input appeared to have compensated the nutrient constraints to phytoplankton during drought. The N:P stoichiometry of deposition and that of lake water indicated that, although there was a seasonal switchover to N- or P-limitation, phytoplankton were mainly co-limited by N and P due probably to the synergistic effects of combined N + P enrichment in the pelagic zone of the lake. Periphyton standing crop showed inverse relationship with phytoplankton and lake DOC. The study indicated that enhanced phytoplankton development and terrestrial DOC flushing in response to atmospheric nutrient input attenuated light penetration to constrain algal periphyton. We suggests that data on these issues may be considered in developing aquatic ecosystem models to establish future links between changing air–water–land interactions and associated shifts in lake ecosystem functioning for more accurately predicting climate change drivers and designing integrated lake basin management strategies.     

Lake‐type‐specific seasonal patterns of nutrient limitation in German lakes,with target nitrogen and phosphorus concentrations for good ecological status

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