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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]