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1.
Nutrient enrichment bioassays, in conjunction with sampling and analysis of surface water chemistry, were conducted in freshwater lakes (kettle ponds) of Cape Cod National Seashore (Massachusetts, USA) to ascertain the importance of nitrogen (N) and phosphorus (P) in regulating the growth of periphyton. Arrays of nutrient diffusing substrata (NDS) were suspended 0.5 m below the water surface in a total of 12 ponds in July and August 2005. Algal biomass developing on each NDS after ~3 weeks of exposure in each month was assessed by quantifying chlorophyll a + phaeophyton pigments. In both July and August, strong responses to N + P and N enrichments were observed in the majority of ponds, while P had no stimulatory effect. These responses correspond well with low atomic ratios (1–18) of dissolved inorganic nitrogen (DIN) to total phosphorus (TP) in ambient surface waters. The results suggest that conditions in the kettle ponds develop whereby nitrogen is the primary limiting nutrient to periphyton growth. While this may be a seasonal phenomenon, it has implications for nutrient management in individual ponds and within the larger watershed.  相似文献   

2.
1. We examined the contribution of algal cells to periphytic organic carbon and assessed the effects of variable biomass composition on the carbon : phosphorus (C : P) ratio of periphyton. We compiled more than 5000 published and unpublished observations of periphytic carbon : chlorophyll a (C : Chl) ratios, an index of algal prevalence, from a variety of substrata collected from lake and low‐salinity coastal habitats. In addition, we converted estimates of algal biovolume into algal C to obtain an independent measure of cellular algal carbon in periphyton. This information was used in a model relating periphyton C : P ratio to algal cellular carbon, the algal C : P ratio, and the C : P ratio of non‐algal organic matter in periphyton. 2. The mean C : Chl ratio of periphyton (405) was relatively high with values in >25% of the samples exceeding 500. On average, 8.4% of total periphyton C was accounted for by C in algal cells. Only 15% of samples were found to have more than 15% periphyton C in cellular algal carbon. Our model showed a nonlinear relationship between periphytic C : P ratios and the C : P ratio of algal cells in the periphyton when non‐algal organic matter was present. However, even at relatively low cellular algal C (<10% of total C), algal C : P ratios can strongly affect the C : P ratio of periphyton as a whole (i.e. algal cells plus other organic matter). 3. The high C : Chl ratios and the low biovolume‐derived algal C of periphyton samples in our data set indicate that algal cells are typically a minor component of organic carbon in periphyton, However, this minor contribution would not preclude algal cellular stoichiometry from notably influencing periphyton C : P ratios.  相似文献   

3.
The potential of periphyton for phosphorus removal from lakes has been investigated using a novel method involving polypropylene (PP) substrate carriers submerged in the pelagial. The study area Lake 'Fühlinger See' in Cologne (Germany) is a complex of mesoeutrophic gravel pit lakes. The whole site is intensively used as a recreation area. Visitors are thought to be the most important single contributors to lake eutrophication. Carriers were exposed at different depths (2, 3.5, 5 m), for different time intervals (1–8 months) and from March to November PP-sheets were readily colonised by periphyton and a biofilm consisting mainly of benthic diatoms developed. Seasonal variability of periphyton on substrates was observed since filamentous green algae colonised the artificial substrates mainly between July and November. Chlorophyll a content of periphyton on the PP-fleece was up to 240-fold higher than chlorophyll a concentrations in the same volume in the epilimnion. Up to around 100 mg of total phosphorus per m2 PP-fleece was bound and can be eliminated from the lake by removal of the substrate carriers together with the periphyton after four months of exposure. Though large-scale validations are needed, this method may be applicable as a technique to harvest phosphorus from the water column in larger-scale settings.  相似文献   

4.
A central premise of ecological stoichiometry is that consumers maintain relatively fixed elemental composition in their bodies, a process known as elemental homoeostasis. Although nutrient enrichment is a ubiquitous problem facing many freshwater lakes around the world, intraspecific variation in elemental composition of freshwater invertebrates and its relationship with nutrient loading have not been well addressed. Here, we examined carbon:nitrogen:phosphorus (C:N:P) stoichiometry of two widely distributed molluscs, Corbicula fluminea and Bellamya aeruginosa, from several subtropical shallow lakes across a nutrient gradient. Our results showed that these two species exhibited substantial natural intraspecific variation in tissue stoichiometry which can reach or even exceed the values among different freshwater taxa investigated before. Our results suggest that tissue P content presents the greatest variations, followed by N content, and lowest in C content. Tissue P content ranged about three-fold (0.56%–1.65%) and five-fold (0.41%–2.28%) for B. aeruginosa and C. fluminea, respectively. Correspondingly, N content ranged from 5.16% to 12.06% and from 6.47 to 11.36%, respectively. Tissue %P, C:P and N:P ratios were strongly correlated with PO43−-P and/or chlorophyll-a in the water column. Tissue N and P contents of B. aeruginosa and P content of C. fluminea increased with increasing lake trophic levels (mesotrophic to eutrophic to hypertrophic). These results suggest that the two molluscs can adjust their tissue stoichiometry in relation to nutrient enrichment. Relaxing the assumption of strict homeostasis may help them cope with potential stoichiometric constraints. The results provide additional clues to why these two species are successful invaders and widely distributed.  相似文献   

5.
Phosphorus enrichment of stream ecosystems generally increases primary production in the benthos, but the consequences of eutrophication for the nutritional quality of periphyton for grazers are less clear. On short timescales, high phosphorus inputs may lead to reduced C:P ratios and high essential fatty acid contents of periphyton, which are both considered important determinants of food quality for grazers. However, nutrient enrichment may alter the taxonomic composition of periphyton and favor the growth of less palatable algal taxa. In this study, periphyton was grown under a gradient of dissolved phosphorus availability from 5 to 100 µg P · L−1, to investigate eutrophication effects on periphyton taxonomy, C:N:P stoichiometry, and fatty acid composition. After 1 month, periphyton grown under oligotrophic conditions was mainly composed of diatoms (~86%). With increasing phosphorus availability, diatoms were gradually outcompeted by chlorophytes and cyanobacteria, which were the predominant taxon under eutrophic conditions. Unexpectedly, periphyton C:P ratios increased with greater phosphorus supply, from ~280 under oligotrophic conditions up to ~790 at 100 µg · L−1, reflecting a tendency of chlorophytes and cyanobacteria to produce more biomass per unit of assimilated phosphorus compared to diatoms. Periphyton content of essential polyunsaturated fatty acids relative to biomass followed a unimodal relationship with phosphorus availability and peaked at intermediate phosphorus levels, likely as a result of both taxonomic and nutrient effects. Our results demonstrate that phosphorus-driven eutrophication of freshwater ecosystems may worsen periphyton nutritional quality due to taxonomic sorting, which may further lead to lower growth and reproduction of herbivores.  相似文献   

6.
Herbivores can have both direct (consumptive) and indirect (nutrient‐mediated) effects on primary producer biomass and nutrient stoichiometry. Ecological stoichiometry theory predicts that herbivores of contrasting body stoichiometry will differentially remineralize nutrients, resulting in feedbacks on producer stoichiometry. We experimentally separated direct and indirect effects of aquatic vertebrate grazers on periphyton by manipulating grazer abundance and identity in mesocosms, and using grazer exclusion cages to expose periphyton to recycled nutrients in the absence of direct grazing. In experiment 1, we used a catfish with high body phosphorus (low body N:P), Ancistrus triradiatus, to assess consumptive versus nutrient‐mediated effects of grazer density on periphyton. In experiment 2, we compared the nutrient‐mediated effects of grazing by Ancistrus triradiatus and Rana palmipes, a tadpole with low body phosphorus and high body N:P. In experiment 1, we found that increasing catfish density led to lower biomass and particulate nutrients in periphyton through direct consumptive effects, but that nutrient‐mediated indirect effects enhanced periphyton biomass when grazers were experimentally separated from direct contact with periphyton. As predicted by stoichiometry theory, nutrient recycling by this P‐rich grazer tended to increase algal C:P and N:P (although effects were not statistically significant), while their consumptive effects reduced algal C:P and N:P. In experiment 2, grazer identity had strong effects on dissolved water nutrient concentrations, N recycling (measured with a 15N tracer), and periphyton stoichiometry. In accordance with stoichiometry theory, catfish increased N concentrations and recycling rates leading to higher periphyton N:P, while tadpoles had greater effects on P availability leading to lower periphyton N:P. Our experiments elucidate the importance of both the density and identity of grazers in controlling periphyton biomass and stoichiometry through consumptive and nutrient‐mediated effects, and support the power of ecological stoichiometry theory to predict feedbacks on producer stroichiometry arising from consumer stoichiometry through nutrient recycling.  相似文献   

7.
We analyzed experimentally the relative contribution of phytoplankton and periphyton in two shallow lakes from the Pampa Plain (Argentina) that represent opposite scenarios according to the alternative states hypothesis for shallow lakes: a clear lake with submerged macrophytes, and a turbid lake with high phytoplankton biomass. To study the temporal changes of both microalgal communities under such contrasting conditions, we placed enclosures in the littoral zone of each lake, including natural phytoplankton and artificial substrata, half previously colonized by periphyton until a mature stage and half clean to analyze periphyton colonization. In the clear vegetated shallow lake, periphyton chlorophyll a concentrations were 3–6 times higher than those of the phytoplankton community. In contrast, phytoplankton chlorophyll a concentrations were 76–1,325 times higher than those of periphyton in the turbid lake. Here, under light limitation conditions, the colonization of the periphyton was significantly lower than in the clear lake. Our results indicate that in turbid shallow lakes, the light limitation caused by phytoplankton determines a low periphyton biomass dominated by heterotrophic components. In clear vegetated shallow lakes, where nitrogen limitation probably occurs, periphyton may develop higher biomass, most likely due to their higher efficiency in nutrient recycling.  相似文献   

8.
1. While the balance of light and nutrients is known to influence the food quality of herbivores by altering algal phosphorus and nitrogen content, the combined effects of light and nutrients on fatty acid synthesis in freshwater periphyton are relatively unknown. In this study, we manipulated light and phosphorus concentration in large, flow‐through experimental streams to examine their effects on both elemental stoichiometry and fatty acid content in periphyton. 2. Two levels of phosphorus (4 and 80 μg L?1) and three of light (17, 40, 110 μmol photons m?2 s?1) were applied in a factorial design in two separate experiments. Diatoms dominated periphyton communities in both experiments, comprising >95% of algal biovolume. Periphyton growth in the streams was simultaneously affected by both resources, even at low rates of supply. 3. Periphyton C/P and C/N ratios increased with light augmentation and decreased with phosphorus enrichment, and consistent with the light : nutrient hypothesis (LNH). Light effects were strongest in streams with low phosphorus concentrations. 4. Periphyton fatty acids reflected the dominance of diatoms : palmitic (16 : 0), palmitoleic (16 : 1ω7) and eicosapentanoic (20 : 5ω3) were the principal saturated (SAFA), monounsaturated (MUFA) and polyunsaturated fatty acids (PUFA), respectively. Linoleic (18 : 2ω6) and linolenic (18 : 3ω3) acids, characteristic of chlorophytes and cyanophytes, were rare, comprising <2% of total fatty acids. 5. Periphyton fatty acid profiles were highly sensitive to light and phosphorus. The proportion of fatty acids comprised by SAFA and MUFA increased with light augmentation and decreased with phosphorus enrichment, whereas PUFA decreased with light and increased with phosphorus. Light effects on fatty acid composition were strongest in phosphorus‐poor streams. PUFA declined with increasing light/phosphorus ratios in the streams, whereas ‘energy’ fatty acids (16 : 0 and 16 : 1) increased. The ratio of SAFA/PUFA was strongly and positively correlated with C/P and C/N ratios. SAFA and MUFA, normalised to dry mass, increased two‐ to threefold with increasing light, while PUFA normalised to dry mass was not significantly affected by light. 6. Similarities in the responses of fatty acids and elemental stoichiometry to light and phosphorus treatments suggested that they were influenced by a common mechanism. Both components of food quality appeared to be sensitive to light‐regulated rates of carbon fixation which, when coupled with insufficient supplies of phosphorus, caused diatom cells to store surplus carbon in SAFA, MUFA and other carbon‐rich compounds that diluted both essential fatty acids and mineral nutrients.  相似文献   

9.
10.
11.
1. To examine how the vertical distribution of periphytic biomass and primary production in the upper 0–1 m of the water column changes along an inter‐lake eutrophication gradient, artificial substrata (plastic strips) were introduced into the littoral zones of 13 lakes covering a total phosphorus (TP) summer mean range from 11 to 536 μg L?1. Periphyton was measured in July (after 8 weeks) and September (after 15 weeks) at three water depths (0.1, 0.5 and 0.9 m). 2. Periphyton chlorophyll a concentration and dry weight generally increased with time and the communities became more heterotrophic. Mean periphytic biomass was unimodally related to TP, reaching a peak between 60 and 200 μg L?1. 3. The proportion of diatoms in the periphyton decreased from July to September. A taxonomic shift occurred from dominance (by biovolume) of diatoms and cyanobacteria at low TP to dominance of chlorophytes at intermediate TP and of diatoms (Epithemia sp.) in the two most TP‐rich lakes. 4. The grazer community in most lakes was dominated by chironomid larvae and the total biomass of grazers increased with periphyton biomass. 5. Community respiration (R), maximum light‐saturated photosynthetic rate (Pmax), primary production and the biomass of macrograzers associated with periphyton were more closely related to periphyton biomass than to TP. Biomass‐specific rates of R, Pmax and production declined with increasing biomass. 6. Mean net periphyton production (24 h) was positive in most lakes in July and negative in all lakes in September. Net production was not related to the TP gradient in July, but decreased in September with increasing TP. 7. The results indicate that nutrient concentrations alone are poor predictors of the standing biomass and production of periphyton in shallow lakes. However, because periphyton biomass reaches a peak in the range of phosphorus concentration in which alternative states occur in shallow lakes, recolonisation by submerged macrophytes after nutrient reduction may potentially be suppressed by periphyton growth.  相似文献   

12.
Silver nanoparticles (AgNPs), like almost all nanoparticles, are potentially toxic beyond a certain concentration because the survival of the organism is compromised due to scores of pathophysiological abnormalities past that concentration. However, the mechanism of AgNP toxicity remains undetermined. Instead of applying a toxic dose, we attempted to monitor the effects of AgNPs at a nonlethal concentration on wild type Drosophila melanogaster by exposing them throughout their development. All adult flies raised in AgNP doped food showed that up to 50 mg/L concentration AgNP has no negative influence on median survival; however, these flies appeared uniformly lighter in body color due to the loss of melanin pigments in their cuticle. Additionally, fertility and vertical movement ability were compromised due to AgNP feeding. Determination of the amount of free ionic silver (Ag+) led us to claim that the observed biological effects have resulted from the AgNPs and not from Ag+. Biochemical analysis suggests that the activity of copper dependent enzymes, namely tyrosinase and Cu-Zn superoxide dismutase, are decreased significantly following the consumption of AgNPs, despite the constant level of copper present in the tissue. Consequently, we propose a mechanism whereby consumption of excess AgNPs in association with membrane bound copper transporter proteins cause sequestration of copper, thus creating a condition that resembles copper starvation. This model also explains the cuticular demelanization effect resulting from AgNP since tyrosinase activity is essential for melanin biosynthesis. Finally, we claim that Drosophila, an established genetic model system, can be well utilized for further understanding of the biological effects of nanoparticles.  相似文献   

13.
Pettersson  Kurt  Grust  Karin  Weyhenmeyer  Gesa  Blenckner  Thorsten 《Hydrobiologia》2003,501(1-3):75-81
The effect of submerged macrophytes on interactions among epilimnetic phosphorus, phytoplankton, and heterotrophic bacterioplankton has been acknowledged, but remains poorly understood. Here, we test the hypotheses that the mean summer phytoplankton biomass (chlorophyll a): phosphorus ratios decrease with increased macrophyte cover in a series of nine lakes. Further, we test that both planktonic respiration and bacterioplankton production increase with respect to phytoplankton biomass along the same gradient of increasing macrophyte cover. Increased macrophyte cover was associated with a lower fraction of particulate phosphorus in epilimnia, with total particulate phosphorus declining from over 80% of total phosphorus in a macrophyte free lake to less than 50% in a macrophyte rich lake. Phytoplankton biomass (chlorophyll a) too was lower in macrophyte dominated lakes, despite relatively high levels of total dissolved phosphorus. Planktonic respiration and bacterioplankton production were higher in macrophyte rich lakes than would be expected from phytoplankton biomass alone, pointing to a subsidy of bacterioplankton metabolism by macrophyte beds at the whole lake scale. The results suggest that the classical view of pelagic interactions, which proposes phosphorus determines phytoplankton abundance, which in turn determines bacterial abundance through the production of organic carbon, becomes less relevant as macrophyte cover increases.  相似文献   

14.
1. We investigated the effects of dissolved organic matter (DOM) and ultraviolet‐B (UVB) radiation on periphyton during a 30‐day experiment in grazer‐free, outdoor artificial streams. We established high [10–12 mg carbon (C) L−1] and low (3–5 mg C L−1) concentrations of DOM in artificial streams exposed to or shielded from ambient UVB radiation. Periphyton was sampled weekly for ash‐free dry mass (AFDM), chlorophyll (chl) a , algal biovolume, elemental composition [C, nitrogen (N) and phosphorus (P)], and algal taxonomic composition. 2. Regardless of the UVB environment, increased DOM concentration caused greater periphyton AFDM, chl a and total C content during the experiment. Increased DOM also significantly increased periphyton C : P and N : P (but not C : N) ratios throughout the experiment. Algal taxonomic composition was strongly affected by elevated stream DOM concentrations; some algal taxa increased and some decreased in biomass and prevalence in artificial streams receiving DOM additions. UVB removal, on the other hand, did not strongly affect periphyton biomass, elemental composition or algal taxonomic composition for most of the experiment. 3. Our results show strong effects of DOM concentration but few, if any, effects of UVB radiation on periphyton biomass, elemental composition and algal taxonomic composition. The effects of DOM may have resulted from its absorption of UVA radiation, or more likely, its provision of organic C and nutrients to microbial communities. The strong effects of DOM on periphyton biomass and elemental composition indicate that they potentially play a key role in food web dynamics and ecosystem processes in forested streams.  相似文献   

15.
Periphyton stoichiometry can vary substantially as a result of differences in stream nutrient availability. A decrease in the periphyton carbon to phosphorus (C:P) ratio should decrease the demand for new P to be immobilized from stream water, but no studies to our knowledge have explored the relationship between periphyton stoichiometry and net P immobilization and release by periphyton. We sought to model biological P immobilization and release (flux) in streams by measuring periphyton stoichiometry and light availability. We measured P flux to and from intact periphyton on stream cobbles (20–100 mm diameter) in 1 L microcosms incubated with streamwater under variable light conditions. Net P immobilization occurred in 75% of microcosms, net P release occurred in only 5% of microcosms, and 20% of microcosms had neither net immobilization nor net release. When normalized to stream conditions, net P immobilization was highest when light availability was high (<60% canopy attenuation) and the periphyton C:P ratio was also high. In contrast, net P release occurred only when light availability was low (>60% canopy attenuation) and the periphyton C:P ratio was also low. A multiple regression model that included both periphyton stoichiometry and light availability from the growing season only, and the interaction term of these two variables, explained 99% of the variation in daily periphyton P flux observed in the study. These results indicate that in order to predict periphyton P immobilization, periphyton stoichiometry and light availability should be considered together. Furthermore, the results indicate that net P immobilization occurs even in very P-rich periphyton, which can act as a P sink when light availability is high.  相似文献   

16.
We examined how mayfly growth rates and body stoichiometry respond to changing phosphorus (P) content in food. In two experiments, mayfly nymphs were given high or low quantities of food at different carbon:phosphorus (C:P) ratios and their growth was measured. Low food quantity resulted in negative growth rates in both experiments, regardless of food P content. However, under high food availability, mayfly growth was affected by the type of food eaten, with low C:P ratio food producing more rapid growth. In addition, mayfly growth increased somewhat when P-poor food was artificially enriched with inorganic P although this effect was not statistically significant. Mayfly body P content was inversely related to body size but increased in animals fed artificially P-enriched food. A model was constructed to simulate mass balance constraints on mayfly growth imposed by the relative supply of two elements (C and P) in food. The model shows that mayfly growth should be limited by food P content at moderately low C:P ratios ( c . 120, by mass). Given high C:P ratios (mean c.  270, by mass) in periphyton from oligotrophic boreal lakes, our experimental and theoretical results indicate that stoichiometric constraints are important factors affecting benthic food webs in lakes from the Canadian Shield and perhaps in other systems with similarly high C:P ratios in periphyton.  相似文献   

17.
Larned  Scott T.  Santos  Scott R. 《Hydrobiologia》2000,432(1-3):101-111
To date, most studies of light- and nutrient-limited primary productivity in forested streams have been carried out in deciduous forests of temperate, continental regions. Conceptual models of light and nutrient limitation have been developed from these studies, but their restricted geographic range reduces the generality of such models. Unlike temperate continental streams, streams on tropical high islands are characterized by flashy, unpredictable discharge and riparian canopies that do not vary seasonally. These contrasting conditions suggest that patterns of light and nutrient limitation in tropical streams may differ from those in temperate streams. The effects of light, and nitrogen and phosphorus availability on periphyton accrual (measured as chlorophyll a per unit area) were investigated using field experiments in 4 low-order streams on the island of Oahu, Hawaii. Levels of chlorophyll a in partially-shaded stream pools were significantly greater than in heavily-shaded pools, and nutrient-enrichment increased the level of chlorophyll a in partially-shaded pools but not in heavily-shaded pools. In each stream, phosphate enrichment resulted in an increase in the level of chlorophyll a, but nitrate enrichment had no effect. Spates following rainstorms occur frequently in these streams, and may increase periphyton productivity by increasing the flux of nutrients to algal cells. However, differences in inorganic nitrogen and phosphorus concentrations measured during spates and baseflow were small, and during some spates, concentrations of these two nutrients declined relative to baseflow concentrations. These observations suggest that phosphorus limitation was not alleviated by spates.  相似文献   

18.
Arnegard  Matthew E.  McCormick  Paul V.  Cairns  John 《Hydrobiologia》1998,385(1-3):163-170
Chemical-diffusing substrates were designed to allow delivery of toxicants to mature periphyton communities under natural conditions without contaminating the surrounding environment. Artificial stream validation studies were conducted in which the effects of substrate-released copper (Cu) on periphyton communities were compared to those generated in a more conventional manner (via water column additions). Effects of copper on the following community parameters were assessed: total community biomass (measured as ash-free dry mass), relative chlorophyll a (chl a and adenosine triphosphate contents, and relative biomass of heterotrophic bacteria. Exposure of more laboratory periphyton communities to substrate-released Cu generated dose-response relationships and recovery models that were indistinguishable from those generated by the conventional route of exposure. The results of this study demonstrate the utility of chemical-diffusing substrates in field validations of laboratory toxicity tests and in investigations of the effects of stress history on periphyton tolerance to toxicants. This revised version was published online in September 2006 with corrections to the Cover Date.  相似文献   

19.
Freshwater gastropods are widespread and common members of benthic communities that interact with other species and conspecifics. Anthropogenic activities are increasing the presence of chemical contaminants in aquatic systems, which have the potential to disrupt species interactions through acute toxic effects and low-exposure chronic effects on vital rates and behaviors of organisms. We determined the effects of the commonly used manufactured nanomaterial, silver nanoparticles (AgNPs), on the survival, growth, reproduction, and behaviors of a common pulmonate gastropod, Physa acuta. Gastropod survival decreased in higher concentrations of AgNP (LC50 = 2.18 μg/l), but was enhanced when experimental containers included sediment (LC50 > 10 μg/l). Chronic exposures resulted in growth rates and size at first reproduction of snails declining in only the highest exposure concentration of 1 μg/l AgNP. Physa egg production was reduced by 50% when chronically exposed to 0.01 μg/l or greater AgNP. Physa crawled more rapidly when exposed to 0.01 μg/l AgNP and greater, indicating a stress response to higher AgNP concentrations. Physa exposed to 1 μg/l AgNP used near-surface habitats in a similar manner to those exposed to the threat of crayfish predation, indicating that the stress response to AgNP is similar in magnitude and direction to the threat of a predator. The sublethal effects reported here suggest that low but environmentally relevant concentrations of AgNP are likely to affect gastropod populations in many ways, potentially leading to measurable effects on communities and ecosystems.  相似文献   

20.
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