Stream Periphyton Research Articles

Terrestrial-aquatic linkage in stream food webs along a forest chronosequence: multi-isotopic evidence.

Long-term monitoring of ecosystem succession provides baseline data for conservation and management, as well as for understanding the dynamics of underlying biogeochemical processes. We examined the effects of deforestation and subsequent afforestation of a riparian forest of Japanese cedar (Cryptomeria japonica) on stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) and natural abundances of radiocarbon (Δ¹⁴C) in stream biota in the Mt. Gomadan Experimental Forest and the Wakayama Forest Research Station, Kyoto University, central Japan. Macroinvertebrates, periphytic algae attached to rock surfaces (periphyton), and leaf litter of terrestrial plants were collected from six headwater streams with similar climate, topography, and bedrock geology, except for the stand ages of riparian forests (from 3 to 49 yr old in five stands and > 90 yr old in one reference stand). Light intensity and δ¹³C values of both periphyton and macroinvertebrates decreased synchronously with forest age in winter. A Bayesian mixing model indicates that periphyton contributions to the stream food webs are maximized in 23-yr-old forests. Except for grazers, most macroinvertebrates showed Δ¹⁴C values similar to those of terrestrial leaf litter, reflecting the influence of modern atmospheric CO₂ Δ¹⁴C values. On the other hand, the Δ¹⁴C values of both periphyton and grazers (i.e., aquatic primary consumers) were significantly lower than that of modern atmospheric CO₂, and were lowest in 23-yr-old forest stands. Previous studies show that root biomass of C. japonica peaks at 15-30 yr after planting. These evidences suggest that soil CO₂ released by root respiration and dispersed by groundwater weathers carbonate substrata, and that dissolved inorganic carbon (DIC) with low Δ¹⁴C is incorporated into stream periphyton and some macroinvertebrates. The ecological response in the studied streams to clear-cutting and replanting of Japanese cedar is much slower (~20 yr) than the chemical response (< 5 yr). More than 50 yr is required for the food web structure to completely recover from clear-cutting. The ecological delay is attributed to several biogeochemical factors, the understanding of which is critical to integrated management of forest-stream continuum and the prediction of ecosystem resilience in response to environmental change.

Effects of light and nutrients on periphyton and the fatty acid composition and somatic growth of invertebrate grazers in subtropical streams.

Algal polyunsaturated fatty acids (PUFA), essential for somatic growth and reproduction of aquatic animals, are influenced by ambient environmental conditions, including light and nutrients. Few studies have addressed the extent to which changes in algal PUFA can influence stream herbivore PUFA profiles and the implications for stream food webs. We manipulated subtropical stream periphyton by applying two light levels (open and shaded canopy) and two nutrient regimes (ambient and enriched) to investigate the response of PUFA and somatic growth in stream herbivores. After 6 weeks, the relative content of periphyton PUFA (%) changed distinctly and differed among treatments. Periphyton in the control treatment with open canopy showed a decline in eicosapentaenoic acid (EPA) relative to initial conditions, whereas shading increased EPA and total highly unsaturated FA (HUFA), but decreased α-linolenic acid (ALA), linoleic acid and total C18 PUFA. The interaction of open canopy and added nutrients increased periphyton ALA compared with initial conditions, while the combined effects of shading and added nutrients led to greater total HUFA. FA similarity between stream grazers (the mayfly Austrophlebioides and caddisfly Helicopsyche) and periphyton increased with periphyton HUFA content. In addition, the growth of large instars of both grazers also increased in response to increased periphyton HUFA%. Our findings show that environmental changes, associated with riparian canopy and nutrients, can lead to changes in periphyton PUFA composition that in turn affect growth and PUFA composition in stream grazers.

Interactive multiple‐stressor effects of the antibiotic monensin, cattle effluent and light on stream periphyton

Summary Streams draining agricultural land are simultaneously exposed to multiple stressors including agricultural antibiotics and effluent from surface water run‐off. The potential interactive effects of these contaminants on stream benthic communities are poorly understood. Using a streamside channel experiment with a full factorial, repeated‐measures design, we assessed the individual and combined effects of the veterinary antibiotic monensin (four concentrations versus control), dairy cattle effluent (added versus control) and differing light levels (reduced versus ambient) on benthic stream periphyton (biomass accrual, community composition and ecological algal guilds). Antibiotic and effluent treatments were applied during a 24‐hour pulse to simulate run‐off from farmland, while light was manipulated throughout the experiment, to mimic the presence or absence of riparian shading. Periphyton taxon richness was decreased by the two highest antibiotic levels 8 days after the pulse. Multivariate community composition (relative abundances of the 15 most common periphyton taxa) was also affected by all three stressors 8 days post‐pulse. However, the periphyton community recovered within 16 days post‐pulse. Interactions among stressors were fairly common, occurring for two of three ecological algal guilds and six of the 15 common taxa. Nevertheless, interactions were generally subtle, without any strong synergistic or antagonistic patterns. Our study shows that the antibiotic monensin can negatively affect stream periphyton communities, but also implies short pulses may not have lasting impacts. The lack of strong synergistic or antagonistic interactions between monensin and light or effluent indicates monensin effects on stream periphyton may be predicted reasonably well based on single‐stressor studies.

Nutrient limitation of algal periphyton in streams along an acid mine drainage gradient.

Metal oxyhydroxide precipitates that form from acid mine drainage (AMD) may indirectly limit periphyton by sorbing nutrients, particularly P. We examined effects of nutrient addition on periphytic algal biomass (chl a), community structure, and carbon and nitrogen content along an AMD gradient. Nutrient diffusing substrata with treatments of +P, +NP and control were placed at seven stream sites. Conductivity and SO4 concentration ranged over an order of magnitude among sites and were used to define the AMD gradient, as they best indicate mine discharge sources of metals that create oxyhydroxide precipitates. Aqueous total phosphorous (TP) ranged from 2 to 23μg·L(-1) and significantly decreased with increasing SO4 . Mean chl a concentrations at sites ranged from 0.2 to 8.1μg·cm(-2) . Across all sites, algal biomass was significantly higher on +NP than control treatments (Co), and significantly increased with +NP. The degree of nutrient limitation was determined by the increase in chl a concentration on +NP relative to Co (response ratio), which ranged from 0.6 to 9.7. Response to nutrient addition significantly declined with increasing aqueous TP, and significantly increased with increasing SO4 . Thus, nutrient limitation of algal biomass increased with AMD impact, indicating metal oxyhydroxides associated with AMD likely decreased P availability. Algal species composition was significantly affected by site but not nutrient treatment. Percent carbon content of periphyton onthe Co significantly increased with AMD impact and corresponded to an increase in the relative abundance of Chlorophytes. Changes in periphyton biomass and cellular nutrient content associated with nutrient limitation in AMD streams may affect higher trophic levels.

Stream primary producers relate positively to watershed natural gas measures in north-central Arkansas streams

Construction of unconventional natural gas (UNG) infrastructure (e.g., well pads, pipelines) is an increasingly common anthropogenic stressor that increases potential sediment erosion. Increased sediment inputs into nearby streams may decrease autotrophic processes through burial and scour, or sediment bound nutrients could have a positive effect through alleviating potential nutrient limitations. Ten streams with varying catchment UNG well densities (0–3.6wells/km2) were sampled during winter and spring of 2010 and 2011 to examine relationships between landscape scale disturbances associated with UNG activity and stream periphyton [chlorophyll a (Chl a)] and gross primary production (GPP). Local scale variables including light availability and water column physicochemical variables were measured for each study site. Correlation analyses examined the relationships of autotrophic processes and local scale variables with the landscape scale variables percent pasture land use and UNG metrics (well density and well pad inverse flow path length). Both GPP and Chl a were primarily positively associated with the UNG activity metrics during most sample periods; however, neither landscape variables nor response variables correlated well with local scale factors. These positive correlations do not confirm causation, but they do suggest that it is possible that UNG development can alleviate one or more limiting factors on autotrophic production within these streams. A secondary manipulative study was used to examine the link between nutrient limitation and algal growth across a gradient of streams impacted by natural gas activity. Nitrogen limitation was common among minimally impacted stream reaches and was alleviated in streams with high UNG activity. These data provide evidence that UNG may stimulate the primary production of Fayetteville shale streams via alleviation of N-limitation. Restricting UNG activities from the riparian zone along with better enforcement of best management practices should help reduce these possible impacts of UNG activities on stream autotrophic processes.

Altitudinal pattern of stream periphyton biomass in tributaries of the Lancang–Mekong River: An indicator of anthropogenic impact?

As the primary producer in streams, periphyton plays an important ecological role in nutrient cycling processes, and has been widely used as an indicator for stream assessment and management. This study examined altitudinal patterns of stream periphyton biomass and their relations to environmental variables in three tributaries of the Lancang–Mekong River. Total dissolved solids, conductivity, and water temperature related negatively with elevation in the three streams. Periphyton biomass showed distinct responses to environmental variables in three streams, and no environmental variables had significant relationship with periphyton biomass in all streams. Altitudinal patterns of periphyton biomass in three streams were uniform: it had a positive relation with elevation, showing a consistent trend with the anthropogenic impact. The altitudinal pattern of stream periphyton biomass could be an ideal indicator to assess stream ecological condition, and monitor anthropogenic impact on mountain streams in the Lancang–Mekong River system.

Silver Nanoparticle Effects on Stream Periphyton During Short-Term Exposures

Silver nanoparticles (AgNP) are increasingly used as antimicrobials in consumer products. Subsequently released into aquatic environments, they are likely to come in contact with microbial communities like periphyton, which plays a key role as a primary producer in stream ecosystems. At present, however, very little is known about the effects of nanoparticles on processes mediated by periphyton communities. We assessed the effects of citrate-coated silver nanoparticles and silver ions (dosed as AgNO3) on five functional end points reflecting community and ecosystem-level processes in periphyton: photosynthetic yield, respiration potential, and the activity of three extracellular enzymes. After 2 h of exposure in experimental microcosms, AgNP and AgNO3 inhibited respiration and photosynthesis of periphyton and the activities of two of the three extracellular enzymes. Addition of a chelating ligand that complexes free silver ions indicated that, in most cases, toxicity of AgNP suspensions was caused by Ag(I) dissolved from the particles. However, these suspensions inhibited one of the extracellular enzymes (leucine aminopeptidase), pointing to a specific nanoparticle effect independent of the dissolved Ag(I). Thus, our results show that both silver nanoparticles and silver ions have potential to disrupt basic metabolic functions and enzymatic resource acquisition of stream periphyton.

Light increases energy transfer efficiency in a boreal stream.

Periphyton communities of a boreal stream were exposed to different light and nutrient levels to estimate energy transfer efficiency from primary to secondary producers using labeling with inorganic 13C. In a one-day field experiment, periphyton grown in fast-flow conditions and dominated by opportunistic green algae were exposed to light levels corresponding to sub-saturating (forest shade) and saturating (open stream section) irradiances, and to N and P nutrient additions. In a two-week laboratory experiment, periphyton grown in low-flow conditions and dominated by slowly growing diatoms were incubated under two sub-saturating light and nutrient enrichment levels as well as grazed and non-grazed conditions. Light had significant positive effect on 13C uptake by periphyton. In the field experiment, P addition had a positive effect on 13C uptake but only at sub-saturating light levels, whereas in the laboratory experiment nutrient additions had no effect on the periphyton biomass, 13C uptake, biovolume and community composition. In the laboratory experiment, the grazer (caddisfly) effect on periphyton biomass specific 13C uptake and nutrient content was much stronger than the effects of light and nutrients. In particular, grazers significantly reduced periphyton biomass and increased biomass specific 13C uptake and C:nutrient ratios. The energy transfer efficiency, estimated as a ratio between 13C uptake by caddisfly and periphyton, was positively affected by light conditions, whereas the nutrient effect was not significant. We suggest that the observed effects on energy transfer were related to the increased diet contribution of highly palatable green algae, stimulated by higher light levels. Also, high heterotrophic microbial activity under low light levels would facilitate energy loss through respiration and decrease overall trophic transfer efficiency. These findings suggest that even a small increase in light intensity could result in community-wide effects on periphyton in boreal streams, with a subsequent increase in energy transfer and system productivity.

Seasonal patterns in stream periphyton fatty acids and community benthic algal composition in six high-quality headwater streams

Fatty acids are integral components of periphyton and differ among algal taxa. We examined seasonal patterns in periphyton fatty acids in six minimally disturbed headwater streams in Pennsylva- nia's Appalachian Mountains, USA. Environmental data and periphyton were collected across four seasons for fatty acid and algal taxa content. Non-metric multidimensional scaling ordination suggested signif- icant seasonal differences in fatty acids; an ordination on algal composition revealed similar seasonal pat- terns, but with slightly weaker separation of summer and fall. Summer and fall fatty acid profiles were driven by temperature, overstory cover, and conduc- tivity and winter profiles by measures of stream size. Ordination on algal composition suggested that sum- mer and fall communities were driven by overstory and temperature, whereas winter communities were driven by velocity. The physiologically important fatty acid 18:3x6 was highest in summer and fall. Winter samples had the highest 20:3x3. Six saturated fatty acids differed among the seasons. Periphyton fatty acids profiles appeared to reflect benthic algal species composition. This suggests that periphyton fatty acid composition can be useful in characterizing basal food resources and stream water quality.

Climate warming and agricultural stressors interact to determine stream periphyton community composition.

Lack of knowledge about how the various drivers of global climate change will interact with multiple stressors already affecting ecosystems is the basis for great uncertainty in projections of future biological change. Despite concerns about the impacts of changes in land use, eutrophication and climate warming in running waters, the interactive effects of these stressors on stream periphyton are largely unknown. We manipulated nutrients (simulating agricultural runoff), deposited fine sediment (simulating agricultural erosion) (two levels each) and water temperature (eight levels, 0-6°C above ambient) simultaneously in 128 streamside mesocosms. Our aim was to determine the individual and combined effects of the three stressors on the algal and bacterial constituents of the periphyton. All three stressors had pervasive individual effects, but in combination frequently produced synergisms at the population level and antagonisms at the community level. Depending on sediment and nutrient conditions, the effect of raised temperature frequently produced contrasting response patterns, with stronger or opposing effects when one or both stressors were augmented. Thus, warming tended to interact negatively with nutrients or sediment by weakening or reversing positive temperature effects or strengthening negative ones. Five classes of algal growth morphology were all affected in complex ways by raised temperature, suggesting that these measures may prove unreliable in biomonitoring programs in a warming climate. The evenness and diversity of the most abundant bacterial taxa increased with temperature at ambient but not with enriched nutrient levels, indicating that warming coupled with nutrient limitation may lead to a more evenly distributed bacterial community as temperatures rise. Freshwater management decisions that seek to avoid or mitigate the negative effects of agricultural land use on stream periphyton should be informed by knowledge of the interactive effects of multiple stressors in a warming climate.

Assessment of periphyton, aquatic macrophytes, benthic communities, and physical habitat in midwestern United States streams coinciding with varying historical concentrations of atrazine

The objectives of this pilot study were to: (1) characterize periphyton and benthic communities using standard collection methods in six Midwest watersheds with varying historical levels of atrazine (low range, medium range and upper range); (2) qualitatively assess presence of aquatic vascular plants at each site; (3) assess and compare physical habitat at each study site in order to evaluate how physical habitat structure may influence the biological communities and (4) analyze the periphyton and benthic macroinvertebrate community data (i.e., series of metrics) among sites to evaluate possible differences or similarities among sites with different historical atrazine exposures. Five of the eight physical habitat metrics (including total physical habitat score) were different among the six study sites. There appeared to be no substantial difference in the structure of periphtyon communities at the six Midwest sites based on 9 of 12 metrics. For the three metrics that showed differences among sites—percentage of sensitive diatoms, percent Achnanches minutissima and percent motile diatoms – there was no consistent pattern with previous degrees of atrazine exposure and the scoring of these metrics. There were also no statistical differences in aquatic macrophyte spatial coverage among the six study areas. Thus, based on the spatially and temporally limited periphyton and aquatic macrophyte data, varying historical atrazine exposure was not associated with impact on resident plant communities (the target receptor group for atrazine). All 10 benthic community metrics showed significant differences among the six Midwest sites. Although no consistent pattern existed with varying historic levels of atrazine, benthic communities at one site with lower historical levels of atrazine were of higher quality than the other five sites. However, this one site also had a higher quality habitat compared to the other sites which was most likely the reason for this benthic condition.

The response of stream periphyton toPacific salmon: using a model to understand the role of environmental context

SummaryIn stream ecosystems,Pacific salmon deliver subsidies of marine‐derived nutrients and disturb the stream bed during spawning. The net effect of this nutrient subsidy and physical disturbance on biological communities can be hard to predict and is likely to be mediated by environmental conditions. For periphyton, empirical studies have revealed that the magnitude and direction of the response to salmon varies from one location to the next. Salmon appear to increase periphyton biomass and/or production in some contexts (a positive response), but decrease them in others (a negative response).To reconcile these seemingly conflicting results, we constructed a system dynamics model that links periphyton biomass and production to salmon spawning. We used this model to explore how environmental conditions influence the periphyton response to salmon.Our simulations suggest that the periphyton response to salmon is strongly mediated by both background nutrient concentrations and the proportion of the stream bed suitable for spawning. Positive periphyton responses occurred when both background nutrient concentrations were low (nutrient limiting conditions) and when little of the stream bed was suitable for spawning (because the substratum is too coarse). In contrast, negative responses occurred when nutrient concentrations were higher or a larger proportion of the bed was suitable for spawning.Although periphyton biomass generally remained above or below background conditions for several months following spawning, periphyton production returned quickly to background values shortly afterwards. As a result, based upon our simulations, salmon did not greatly increase or decrease overall annual periphyton production. This suggests that any increase in production by fish or invertebrates in response to returning salmon is more likely to occur via direct consumption of salmon carcasses and/or eggs, rather than the indirect effects of greater periphyton production.Overall, our simulations suggest that environmental factors need to be taken into account when considering the effects of spawning salmon on aquatic ecosystems. Our model offers researchers a framework for testing periphyton response to salmon across a range of conditions, which can be used to generate hypotheses, plan field experiments and guide data collection.

Effects of a chronic lower range of triclosan exposure on a stream mesocosm community

Triclosan (5-chloro-2-(2,4-dichlorophenoxy)phenol) is an antimicrobial found in consumer soaps and toothpaste. It is in treated wastewater effluents at low parts-per-billion concentrations, representing a potentially chronic exposure condition for biota inhabiting receiving streams. For the present study, a naturally colonized benthos was created using flow-through indoor mesocosms; then, the benthic communities were dosed to achieve different in-stream triclosan concentrations (control, 0.1 μg/L, 0.5 μg/L, 1.0 μg/L, 5.0 μg/L, and 10 μg/L) for 56 d. Water quality parameters and endpoints from bacteria to macroinvertebrates, as well as interacting abiotic components, were measured. Effects of triclosan on specific microbial endpoints were observed at all doses, including an effect on litter decomposition dynamics at doses of 1.0 μg/L and higher. Resistance of periphytic bacteria to triclosan significantly increased at doses of 0.5 μg/L and above. By the end of dosing, the antimicrobial appeared to stimulate the stream periphyton at the 3 lowest doses, while the 2 highest doses exhibited decreased stocks of periphyton, including significantly lower bacteria cell densities and cyanobacteria abundance compared with the control. Other than an effect on benthic ostracods, the changes that occurred in the periphyton did not translate to significant change in the colonizing nematodes, the macroinvertebrate community as a whole, or other measurements of stream function. The results shed light on the role a low, chronic exposure to triclosan may play in effluent-dominated streams.

Quantifying the direct and indirect effects of flow‐related disturbance on stream fish assemblages

Summary Flood‐related disturbances are predicted to be seriously altered by climate change effects, and this will have strong implications for stream communities. Predicting how and why community structure responds to changes in disturbance regimes will require measures of disturbance that are closely linked to community variability. A range of disturbance measures have been tested for their ability to explain patterns in stream periphyton and invertebrate assemblages, but assessments of fish have largely focussed on flow measures as predictors. Consequently, the mechanisms driving fish assemblage responses to disturbance are poorly known. Stream disturbance was quantified using seven measures, three based on disturbance of the streambed (assessed using painted tracer particles), three associated with variation in stream discharge and one from a subjective measure of stream channel stability. Twenty streams were sampled on five occasions to evaluate which disturbance measure explained the most variation in fish assemblages and also to quantify the influence of disturbance on spatial and temporal assemblage variability. To determine whether disturbance affected fish directly or via its effect on benthic food supply (i.e. stream invertebrates), a one‐off survey of 52 streams was conducted. Path analysis on data from the 20 streams indicated that fish biomass responded most strongly to flow‐mediated bed movement, not flow variability. Consequently, bed‐movement measures were better predictors of spatial and temporal variability in fish biomass and assemblage structure than purely hydrological measures. Furthermore, variations in fish biomass and assemblage structure were higher in more disturbed streams. A second path analysis conducted on data from the 52 stream survey compared the effects of physical habitat and food‐related components of bed disturbance on fish biomass. It indicated that both had a similar influence on disturbance‐related reductions in fish biomass. These results indicate that variation in fish assemblages can best be understood by measuring or predicting bed movement, because bed disturbance is likely to encompass more effectively major factors that influence stream fish, such as habitat availability and food supply. A shift in disturbance regime is likely to change the biomass of fish a habitat can support and fish assemblage structure.

Understanding the combined influence of fine sediment and glyphosate herbicide on stream periphyton communities

Pesticides and deposited fine sediment have independently been associated with changes in relative abundance and species richness in aquatic ecosystems, but the interplay between these two stressors in agricultural streams is poorly understood. A 28-day experiment in outdoor stream mesocosms examined the effects of four levels each of fine sediment coverage (0, 25, 75, 100%) and glyphosate-based herbicide concentration (0, 50, 200, 370 μg/L) on periphyton communities (algae and bacteria) in a fully factorial, repeated-measures design. Our aims were to determine whether (i) increased levels of sediment and glyphosate had individual and/or additive effects, (ii) increased sediment reduced the toxicity of glyphosate (antagonistic multiple stressor interaction), or (iii) sediment-adsorbed glyphosate prolonged the effects of exposure (synergistic interaction). We also assigned all algal taxa to three ecological guilds (low-profile, high-profile and motile growth forms) and separately determined their responses to the treatments. As individual stressors, sediment addition affected all algal community-level metrics, whereas glyphosate addition only affected algal community evenness. Bacterial taxon richness was unaffected by either stressor. In combination, however, significant overall sediment by glyphosate interactions were detected, demonstrating synergistic (algal evenness, high-profile and motile guilds) or antagonistic effects (low-profile guild). Our experiment underscores the importance of considering both structural and functional indicators, including algal guild representation, when assessing the mechanisms by which periphyton communities respond to multiple stressors.

Longitudinal patterns of periphyton biomass in Qinghai–Tibetan Plateau streams: An indicator of pasture degradation?

Periphyton communities play a key role in stream ecosystems, and are usually used as an indicator for long-term biological monitoring, management and evaluation. This study examined the relationships between environmental variables and stream periphyton biomass, investigated the longitudinal patterns of periphyton biomass, and explored the relationships between these longitudinal patterns and pasture degradations in Qinghai–Tibetan Plateau. Isolated environmental variables did not have significant impacts on periphyton biomass and did not clearly respond to the influence of pasture degradation. The longitudinal patterns of stream periphyton biomass were consistent with pasture degradation trends. The more serious the pasture degradation, the lower is the periphyton biomass. Longitudinal gradient is a comprehensive variable, which represents a number of space-related environmental factors. In Qinghai–Tibetan Plateau, the pasture ecosystems are fragile and have been severely degraded. The longitudinal patterns of periphyton biomass were closely related to ecological degradation and can be used as an indicator to monitor and evaluate pasture degradation in Qinghai–Tibetan Plateau.

Elevated light and nutrients alter the nutritional quality of stream periphyton

Summary The biochemical composition of primary food resources may affect secondary production, growth, reproduction and other physiological responses in consumers and may be an important driver of food‐web dynamics. Changes in land use, riparian clearing and non‐point nutrient inputs to streams have the potential to alter the biochemical composition of periphyton, and characterising this relationship may be critical to understand the processes by which environmental change can affect food webs and ecosystem function. We conducted a manipulative, in situ experiment to examine the effect of light and nutrient availability on stream periphyton biomass, nutrient content, stoichiometry and fatty acid composition. Greater light increased periphyton biomass [chl‐a, ash‐free dry mass (AFDM)], periphyton carbon concentrations and monounsaturated fatty acids (MUFAs), but decreased saturated fatty acids (SAFAs). Greater light availability also increased levels of &lt;20C polyunsaturated fatty acids (PUFAs), but decreased quantities of several long‐chain (20–22 C), highly unsaturated fatty acids (HUFAs). Nutrient (+N, +P) addition had no significant effect on periphyton biomass in the study streams, but did increase periphyton carbon content. For fatty acids, despite non‐significant effects on periphyton biomass, nutrient additions resulted in an increased ratio of SAFA to PUFA, greater concentrations of stearidonic acid (18:4ω3) and near‐significant increases in α‐linolenic acid (18:3ω3). Nutrient additions also resulted in decreased quantities of the physiologically important docosahexaenoic acid (DHA; 22:6ω3) and increased the ratio of omega‐3 to omega‐6 fatty acids. An interaction between nutrient addition and greater light also resulted in decreased percentages of another physiologically important fatty acid, eicosapentaenoic acid (EPA; 20:5ω3). Results from this study suggest that increased light and nutrients, a condition common in urban and agricultural catchments, would increase 18C PUFA intermediates that would be available in periphyton food sources, but decrease the availability of essential 20‐22C HUFAs. If stream consumers are unable to elongate PUFAs into essential HUFAs, there may be important effects on food webs, including shifts by invertebrate consumers from herbivory to omnivory or carnivory, or changes in the species composition of aquatic consumers. Our results demonstrate that fatty acid measures were more sensitive to changes in light and nutrient conditions than more traditional measures of biomass, nutrient content and stoichiometry. We recommend routine measurement of fatty acid content in monitoring the effect of changing environmental conditions in rivers and streams, particularly studies on food quality and ecosystem health.

Rates of Species Accumulation and Taxonomic Diversification during Phototrophic Biofilm Development Are Controlled by both Nutrient Supply and Current Velocity

The accumulation of new and taxonomically diverse species is a marked feature of community development, but the role of the environment in this process is not well understood. To address this problem, we subjected periphyton in laboratory streams to low (10-cm · s(-1)), high (30-cm · s(-1)), and variable (9- to 32-cm · s(-1)) current velocity and low- versus high-nutrient inputs. We examined how current velocity and resource supply constrained (i) the rates of species accumulation, a measure of temporal beta-diversity, and (ii) the rates of diversification of higher taxonomic categories, defined here as the rate of higher taxon richness increase with the increase of species richness. Temporal biofilm dynamics were controlled by a strong nutrient-current interaction. Nutrients accelerated the rates of accumulation of new species, when flow velocity was not too stressful. Species were more taxonomically diverse under variable than under low-flow conditions, indicating that flow heterogeneity increased the niche diversity in the high-nutrient treatments. Conversely, the lower diversification rates under high- than under low-nutrient conditions at low velocity are explained with finer resource partitioning among species, belonging to a limited number of related genera. The overall low rates of diversification in high-current treatments suggest that the ability to withstand current stress was conserved within closely related species. Temporal heterogeneity of disturbance has been shown to promote species richness, but here we further demonstrate that it also affects two other components of biodiversity, i.e., temporal beta-diversity and diversification rate. Therefore, management efforts for preserving the inherent temporal heterogeneity of natural ecosystems will have detectable positive effects on biodiversity.

Consistency and sensitivity of stream periphyton community structural and functional responses to nutrient enrichment

Eutrophication remains one of the foremost impacts of industrialization and population expansion on aquatic ecosystems worldwide. Selecting metrics for assessing the manner in which communities and biogeochemical processes respond to nutrient fertilization is an ongoing management challenge critical both for detecting changes and for monitoring recovery of impaired environments. A key limitation to the selection of response variables is the lack of consistent evaluation of metrics under the same conditions in multiple systems across ecoregions. Here we report the results of nutrient-diffusing agar fertilization experiments conducted simultaneously in 30 streams in two distinct ecoregions of California, USA: the mountainous Sierra Nevada and the coastal chaparral of Santa Barbara county. In each experiment we evaluated algal community shifts across five nutrient delivery rates using multiple response variables at the ecosystem process (respiration and primary production), community (biomass and diversity metrics), functional group (nitrogen fixing, growth form, nutrient adaptation), and taxonomic group (indicator species, genera, families) levels of ecological organization. We used mixed-effect general linear models to quantify the magnitude, sensitivity, and consistency of responses among streams within and across ecoregions to provide an objective assessment of the potential for each variable to describe and detect significant changes in algal community characteristics. Our results indicate that ecosystem- and community-level variables showed significant and consistent nutrient responses among diverse streams and across the two ecoregions, while indicator taxa and functional groups were less likely to respond consistently to nutrient enrichment. We discuss the relevance of our findings to the ongoing development of monitoring and bioassessment strategies for aquatic eutrophication.

Divergent responses of biomass and enzyme activities suggest differential nutrient limitation in stream periphyton

We used in situ nutrient-enrichment experiments at sites in 4 Texas streams spanning a steep nutrient gradient to test the hypothesis that periphyton photoautotrophs and heterotrophs are differentially limited by N and P, respectively. We contrasted the responses of total periphyton biomass (ash-free dry mass [AFDM]), photoautotrophic biomass (chlorophyll a [CHLA]), % CHLA of AFDM [% CHLA]), alkaline phosphatase activity (APA), and nitrogenase activity (NA) to ambient N and P concentrations among sites and among nutrient-diffusing substrata (NDS) (control, N, P, and N+P) within sites. Periphyton biomass (AFDM and CHLA) increased steadily with increasing ambient N and P among sites. Both APA and NA were relatively high on the control treatments at the lowest nutrient site and decreased with increasing ambient N and P until falling to negligible levels at the highest nutrient site. Within sites, P enrichment decreased APA but did not increase % CHLA or CHLA/cm2. P enrichment did increase AFDM/cm2 at one of the low nutrient sites. N enrichment reduced NA and increased % CHLA and CHLA/cm2 without increasing AFDM/cm2. N+P enrichment resulted in the highest % CHLA and CHLA/cm2 at the 3 lowest nutrient sites but did not increase AFDM/cm2. Combined, these findings suggest that photoautotrophs in periphyton communities may be more limited than heterotrophs by N or become more competitive for P under elevated N. The high levels of APA and NA on the controls coupled with minimal difference in AFDM among nutrient treatments at most sites despite increases in % CHLA on N and N+P treatments provide experimental evidence that differential nutrient limitation probably occurs in periphyton communities. Our results support the view that the single-limiting-nutrient paradigm may lead to overly simplistic interpretations of periphyton responses to nutrient enrichment.