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Summary 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. [ABSTRACT FROM AUTHOR]

Light and nutrients play pivotal roles in determining the growth of autotrophs, yet the potential for synergistic interactions between the two resources in algal communities is poorly understood, especially in stream ecosystems. In this study, light and phosphorus were manipulated in large experimental streams to examine resource colimitation and synergy in stream periphyton. Whole-stream metabolism was simultaneously limited by light and phosphorus. Increasing the supply of either light or phosphorus resulted in significant increases in primary production and the transformation of the streams from heterotrophy to autotrophy. Resource-driven changes in periphyton community structure occurred in concert with changes in production. Algal assemblages in highly shaded streams were composed primarily of small diatoms such as Achnanthidium minutissima, whereas larger diatoms such as Melosira varians predominated at higher irradiances. Phosphorus enrichment had relatively little effect on assemblage structure, but it did substantially diminish the abundance of Meridion circulare, a diatom whose mucilaginous colonies were conspicuously abundant in phosphorus-poor, high-light streams. Bacterial biomass declined relative to algal biomass with increases in primary productivity, regardless of whether the increases were caused by light or phosphorus. Synergistic effects on primary production appeared to occur because the availability of one resource facilitated the utilization of the other. Light increased the abundance of large diatoms, which are known to convert high concentrations of nutrients into primary production more effectively than smaller taxa. Phosphorus enrichment led to the replacement of Meridion circulare by non-mucilaginous taxa in phosphorus-enriched streams, and we hypothesize that this change enabled more efficient use of light in photosynthesis. Higher ratios of chlorophyll a : biomass in phosphorus-enriched streams may have also led to more efficient photon capture and higher photosynthetic rates. Synthesis. Our results underscore the potential for resource colimitation, even in habitats where a single resource is as strongly limiting as is light in shaded streams. The capacity of autotrophic communities to respond to more than one limiting resource suggests that prevailing single-resource models of ecosystem productivity are overly simplistic. [ABSTRACT FROM AUTHOR]

1. The light : nutrient hypothesis (LNH) states that algal nutrient content is determined by the balance of light and dissolved nutrients available to algae during growth. Light and phosphorus gradients in both laboratory and natural streams were used to examine the relevance of the LNH to stream periphyton. Controlled gradients of light (12–426 μmol photons m−2 s−1) and dissolved reactive phosphorus (DRP, 3–344 μg L−1) were applied experimentally to large flow-through laboratory streams, and natural variability in canopy cover and discharge from a wastewater treatment facility created gradients of light (0.4–35 mol photons m−2 day−1) and DRP (10–1766 μg L−1) in a natural stream. 2. Periphyton phosphorus content was strongly influenced by the light and DRP gradients, ranging from 1.8 to 10.7 μg mg AFDM−1 in the laboratory streams and from 2.3 to 36.9 μg mg AFDM−1 in the natural stream. Phosphorus content decreased with increasing light and increased with increasing water column phosphorus. The simultaneous effects of light and phosphorus were consistent with the LNH that the balance between light and nutrients determines algal nutrient content. 3. In experiments in the laboratory streams, periphyton phosphorus increased hyperbolically with increasing DRP. Uptake then began levelling off around 50 μg L−1. 4. The relationship between periphyton phosphorus and the light : phosphorus ratio was highly nonlinear in both the laboratory and natural streams, with phosphorus content declining sharply with initial increases in the light : phosphorus ratio, then leveling off at higher values of the ratio. 5. Although light and DRP both affected periphyton phosphorus content, the effects of DRP were much stronger than those of light in both the laboratory and natural streams. DRP explained substantially more of the overall variability in periphyton phosphorus than did light, and light effects were evident only at lower phosphorus concentrations (≤25 μg L−1) in the laboratory streams. These results suggest that light has a significant negative effect on the food quality of grazers in streams only under a limited set of conditions. [ABSTRACT FROM AUTHOR]

The light nutrient hypothesis posits that herbivore growth is increasingly constrained by low food quality as the ratio of light to nutrients increases in aquatic ecosystems. We tested predictions of this hypothesis by examining the effects of large seasonal cycles in light and nutrients on the mineral content of periphyton and the growth rate of a dominant herbivore (the snail Elimia clavaeformis) in two oligotrophic streams. Streambed irradiances in White Oak Creek and Walker Branch (eastern Tennessee, USA) varied dramatically on a seasonal basis due to leaf phenology in the surrounding deciduous forests and seasonal changes in sun angle. Concentrations of dissolved nutrients varied inversely with light, causing light: nitrate and light: phosphate to range almost 100-fold over the course of any individual year. Periphyton nitrogen and phosphorus concentrations were much lower than the concentrations of these elements in snails, and they bottomed out in early spring when streambed irradiances were highest. Snail growth, however, peaked in early spring when light: nutrient ratios were highest and periphyton nutrient concentrations were lowest. Growth was linearly related to primary production (accounting for up to 85% of growth variance in individual years), which in turn was driven by seasonal variation in light. Conceptual models of herbivore growth indicate that growth should initially increase as increasing light levels stimulate primary production, but then level off, and then decrease as the negative effects of decreasing algal nutrient content override the positive effects of increased food production. Our results showed no, evidence of an inflection point where increasing ratios of light to nutrients negatively affected growth. Snail growth in these intensively grazed streams is probably unaffected by periphyton nutrient content because exploitative competition for food reduces growth rates to levels where the demand for nitrogen and phosphorus is small enough to be satisfied by even low levels of these nutrients in periphyton. Competition for limited food resources in habitats where herbivore densities are uncontrolled by predation or other mortality factors should strongly influence the potential for herbivores to be limited by mineral deficits in their food. [ABSTRACT FROM AUTHOR]

1. This study investigated the combined effects of light and phosphorus on the growth and phosphorus content of periphyton. To investigate the potential for colimitation of algal growth by these two resources, diatom-dominated periphyton communities in large flow-through laboratory streams were exposed under controlled conditions to simultaneous gradients of light and phosphorus. 2. Periphyton growth rate was predictably light-limited by the subsaturating irradiances (12–88 μmol photons m−2 s−1) used in this experiment. However, phosphorus concentration also limited growth rate: growth increased hyperbolically with increasing soluble reactive phosphorus (SRP), reaching a threshold of growth saturation between 22 and 82 μg L−1. 3. Periphyton phosphorus content was strongly and nonlinearly related with SRP, reaching a maximum at 82 μg L−1 SRP. Contrary to the Light : Nutrient Hypothesis, periphyton phosphorus content did not decrease with increasing light, even at the lowest concentrations of SRP. Periphyton phosphorus was highly correlated with periphyton growth rate (Spearman's ρ = 0.63, P < 0.005). 4. Multiple regression analysis reinforced evidence of simultaneous light and phosphorus limitation. Both light and periphyton phosphorus content were significant variables in multiple regressions with growth parameters as dependent variables. Light alone accounted for 67% of the variance in periphyton biomass, and the addition of periphyton phosphorus as an additional independent variable increased the total amount of variance explained to 81%. 5. Our results did not support the hypothesis that extra phosphorus is required for photoacclimation to low light levels. Rather, the effect of additional phosphorus may have been to accommodate increased requirements for P-rich ribosomal RNA when growth was stimulated by increased light. The potential colimitation of periphyton growth by phosphorus and light at subsaturating irradiances has important implications in both theoretical and applied aquatic ecology. [ABSTRACT FROM AUTHOR]

1. Streambed light regimes change dramatically when riparian trees gain leaves in spring and lose them in autumn. This study examined the effect of these changes on periphyton photosynthetic characteristics, primary production, and light utilisation efficiency in two eastern Tennessee streams. 2. Photosynthesis–irradiance responses were measured at intervals covering leaf emergence and abscission in spring and autumn. Photosynthetic efficiency (αchl ) increased with declining streambed irradiances during spring leaf emergence, but returned to pre-emergence values after autumn leaf fall. The onset of photosaturation (I k ) displayed the opposite pattern, decreasing during leaf emergence and increasing after leaf fall. Both αchl and I k were closely associated (P < 0.01) with daily integrated streambed irradiance, as were periphyton carotenoids. Internal shading by photoprotective carotenoids is hypothesised to account for lower αchl when streambed irradiances are high. 3. An in situ shading experiment confirmed that the temporal changes observed in periphyton photosynthetic characteristics and carotenoids were primarily the result of changing light levels and not other environmental factors (e.g. nutrients, temperature). 4. Daily chlorophyll-specific primary production (PP chl ) was calculated with P –I models and recorded streambed irradiances. In both streams, PP chl was the highest in early spring when trees were leafless, and then declined markedly as leaves emerged, reaching a minimum in summer. PP chl increased after leaf abscission, but was still lower than it was in early spring, when the sun was higher and daylength was longer. A hyperbolic tangent equation fit to PP chl and daily integrated irradiance (r 2 =0. 85) suggested that primary production was light saturated at 4–8 mol m–2... [ABSTRACT FROM AUTHOR]

Summary 1 This laboratory study examined the effect of a gradient of UV-B radiation (280–320 nm) on photosynthesis and food quality of periphyton, the trophic base of many freshwater benthic communities. Four irradiances of UV-B (0, 0.6, 1.2, and 2.3 W m-2) were delivered by UV-B lamps (313 nm peak irradiance) over a 13-day period in the first experiment and over a 4-h period in the second experiment. These irradiances were roughly equivalent to 0, 1, 2, and 4 times the ambient biologically effective (DNA) midsummer, midday UV-B irradiance in Tennessee. 2 Rates of photosynthesis and photosynthetic pigments were significantly reduced by irradiances greater than ambient during the 13-day experiment, suggesting that food supply rates to grazers would be depressed by increases in current UV-B levels. Effects on community structure were minor, but mean diatom cell size decreased at higher UV-B irradiances. 3 Irradiated periphyton was fed in surplus to juvenile snails (Physella gyrina) in the first experiment as a bioassay for food quality. Snail growth was the same on all four diets, suggesting that UV-B did not affect food quality. Nitrogen and phosphorus content of the periphyton were not affected by UV-B, either. 4 Photosynthesis by low-biomass periphyton in the second experiment was significantly depressed by irradiances above ambient after only 4 h. Photosynthesis by the high biomass periphyton was not significantly affected by UV-B, suggesting that self-shading reduced UV-B effects. [ABSTRACT FROM AUTHOR]

Examines the response of periphyton and grazing snails (Elimia clavaeformis) to summer shade in a Tennessee forest. Effects of light limitation and grazing; Shade adaptation; Snail growth versus irradiance; Snail growth; Lipid accumulation; Photosynthesis-irradiance curves.

Stable isotopes are widely used to infer trophic relationships with little attention paid to temporal variability at the base of the food web. We examined changes in the carbon-stable isotope composition during periphyton development, sampling periphyton that accumulated on ceramic tiles at four stream sites over a 2-month period. Periphyton 13C rose and fell in general concordance with rising and falling biomass at all four sites, resulting in significant correlations between periphyton δ13C and chlorophyll a (Chl a). Mean δ13C values at one site rose from -26% to -20% in 2 weeks, falling back to -24% the next week after a large scouring spate. Periphyton 13C also underwent a smaller, longer-term increase that correlated with a gradual rise in stream temperature. Multiple regression analysis with both Chl a and temperature as independent variables accounted for up to 88% of the temporal variability in δ13C, with Chl a the largest source of variability. Water velocity, measured on each sampling occasion, was unrelated to temporal changes in 13C. Depletion of inorganic carbon within the periphyton matrix is the probable cause of increasing 13C in periphyton as biomass develops. Rising δ13C values during periphyton biomass development suggest the possibility of carbon-limited periphyton growth, even in alkaline waters. The strong link between biomass and periphyton 13C helps explain the large range of δ13C reported for periphyton in streams, where temporal and spatial variability in periphyton biomass are notorious. [ABSTRACT FROM AUTHOR]

Headwater streams are key sites of nutrient and organic matter processing and retention, but little is known about temporal variability in gross primary production (GPP) and ecosystem respiration (ER) rates as a result of the short duration of most metabolism measurements in lotic ecosystems. We examined temporal variability and controls on ecosystem metabolism by measuring daily rates continuously for 2 years in Walker Branch, a first-order deciduous forest stream. Four important scales of temporal variability in ecosystem metabolism rates were identified: (1) seasonal, (2) day-to-day, (3) episodic (storm-related), and (4) inter-annual. Seasonal patterns were largely controlled by the leaf phenology and productivity of the deciduous riparian forest. Walker Branch was strongly net heterotrophic throughout the year with the exception of the open-canopy spring when GPP and ER rates were co-equal. Day-to-day variability in weather conditions influenced light reaching the streambed, resulting in high day-to-day variability in GPP particularly during spring (daily light levels explained 84% of the variance in daily GPP in April). Episodic storms depressed GPP for several days in spring, but increased GPP in autumn by removing leaves shading the streambed. Storms depressed ER initially, but then stimulated ER to 2–3 times pre-storm levels for several days. Walker Branch was strongly net heterotrophic in both years of the study, with annual GPP being similar (488 and 519 g O2 m−2 y−1 or 183 and 195 g C m−2 y−1) but annual ER being higher in 2004 than 2005 (−1,645 vs. −1,292 g O2 m−2 y−1 or −617 and −485 g C m−2 y−1). Inter-annual variability in ecosystem metabolism (assessed by comparing 2004 and 2005 rates with previous measurements) was the result of the storm frequency and timing and the size of the spring macroalgal bloom. Changes in local climate can have substantial impacts on stream ecosystem metabolism rates and ultimately influence the carbon source and sink properties of these important ecosystems. [ABSTRACT FROM AUTHOR]