Factors influencing periphyton growth in agricultural streams of central Illinois

Factors limiting periphyton accrual in east-central Illinois agricultural streams were investigated. Nutrient-diffusing substrata were used to examine periphyton macronutrient limitation in streams in two agricultural watersheds. Substrata consisted of sand-agar mixtures with one of six experimental treatments. Macronutrients included carbon, nitrate, phosphate and combinations of the three. Substrata were collected after a 5 and 9 day period and analyzed for chlorophyll a. None of the treatments were significantly greater than the controls at any of the seven stations, thus we conclude that periphyton in these streams was not nutrient limited. Highest periphyton colonization/growth rates were associated with the smaller upstream reaches, while lower rates occurred in the larger downstream reaches. Multiple regression showed that most of the variance in the rate of chlorophyll a accrual after five days was explained through water temperature and turbidity (r² = 0.91); whereas, stream nitrate and phosphate concentrations accounted for no significant portion of the variance. We conclude that instream primary production in agricultural streams of central Illinois is limited by temperature and light.     

Stream food web response to a salmon carcass analogue addition in two central Idaho, U.S.A. streams

1. Pacific salmon and steelhead once contributed large amounts of marine‐derived carbon, nitrogen and phosphorus to freshwater ecosystems in the Pacific Northwest of the United States of America (California, Oregon, Washington and Idaho). Declines in historically abundant anadromous salmonid populations represent a significant loss of returning nutrients across a large spatial scale. Recently, a manufactured salmon carcass analogue was developed and tested as a safe and effective method of delivering nutrients to freshwater and linked riparian ecosystems where marine‐derived nutrients have been reduced or eliminated. 2. We compared four streams: two reference and two treatment streams using salmon carcass analogue(s) (SCA) as a treatment. Response variables measured included: surface streamwater chemistry; nutrient limitation status; carbon and nitrogen stable isotopes; periphyton chlorophyll a and ash‐free dry mass (AFDM); macroinvertebrate density and biomass; and leaf litter decomposition rates. Within each stream, upstream reference and downstream treatment reaches were sampled 1 year before, during, and 1 year after the addition of SCA. 3. Periphyton chlorophyll a and AFDM and macroinvertebrate biomass were significantly higher in stream reaches treated with SCA. Enriched stable isotope (δ¹⁵N) signatures were observed in periphyton and macroinvertebrate samples collected from treatment reaches in both treatment streams, indicating trophic transfer from SCA to consumers. Densities of Ephemerellidae, Elmidae and Brachycentridae were significantly higher in treatment reaches. Macroinvertebrate community composition and structure, as measured by taxonomic richness and diversity, did not appear to respond significantly to SCA treatment. Leaf breakdown rates were variable among treatment streams: significantly higher in one stream treatment reach but not the other. Salmon carcass analogue treatments had no detectable effect on measured water chemistry variables. 4. Our results suggest that SCA addition successfully increased periphyton and macroinvertebrate biomass with no detectable response in streamwater nutrient concentrations. Correspondingly, no change in nutrient limitation status was detected based on dissolved inorganic nitrogen to soluble reactive phosphorus ratios (DIN/SRP) and nutrient‐diffusing substrata experiments. Salmon carcass analogues appear to increase freshwater productivity. 5. Salmon carcass analogues represent a pathogen‐free nutrient enhancement tool that mimics natural trophic transfer pathways, can be manufactured using recycled fish products, and is easily transported; however, salmon carcass analogues should not be viewed as a replacement for naturally spawning salmon and the important ecological processes they provide.     

Controls on periphyton biomass in heterotrophic streams

1. Headwater streams of the Hubbard Brook Experimental Forest (HBEF) are typically characterised by a periphyton assemblage of low biomass and diversity. However, periphyton blooms have been observed following catchment deforestation experiments and occasionally during the annual spring thaw before canopy leaf‐out. 2. There is pronounced seasonal variation in both nutrient and light availability in HBEF streams. Stream water nitrogen (N) concentrations and light levels are higher before canopy leaf‐out and after leaf senescence and are lower during the growing season. Periphyton accrual rates also change seasonally; they are highest in spring prior to leaf‐out and significantly lower during summer and in autumn. 3. Periphyton biomass rarely responded positively to in‐situ experimental enrichment with nitrogen or phosphorus. In the summer, nutrient enrichment overall had no effect on periphyton biomass, while outside the growing season N enrichment had inhibitory effects on periphyton. 4. Despite these experimental results, surveys of ambient chlorophyll a concentrations in streams across the HBEF demonstrated no relationship between streamwater dissolved inorganic N or P concentrations and benthic chlorophyll a. 5. Our results suggest that HBEF periphyton communities are not closely regulated by nutrient availability, even during periods of high light availability. The inhibitory effects of nutrient enrichment outside the growing season are interesting, but further research is necessary to elucidate the mechanisms driving these responses.     

Quantitative Links Between Pacific Salmon and Stream Periphyton

Species’ impacts on primary production can have strong ecological consequences. In freshwater ecosystems, Pacific salmon (Oncorhynchus spp.) may influence stream periphyton through substrate disturbance during spawning and nutrient subsidies from senescent adults. The shape of relationships between the abundance of spawning salmon and stream periphyton, as well as interactions with environmental variables, are incompletely understood and may differ across the geographic range of salmon. We examined these relationships across 24 sockeye salmon (Oncorhynchus nerka) spawning streams in north-central British Columbia, Canada. The influence of salmon abundance and environmental variables (temperature, light, dissolved nutrients, water velocity, watershed size, and invertebrate grazer abundance) on post-spawning periphyton abundance and nitrogen stable isotope signatures, which can indicate the uptake of salmon nitrogen, was evaluated using linear regression models and Akaike Information Criterion. Periphyton nitrogen stable isotope signatures were best described by a positive log-linear relationship with an upstream salmon abundance metric that includes salmon from earlier years. This suggests the presence of a nutrient legacy. In contrast, periphyton abundance was negatively related to the spawning-year salmon density, which likely results from substrate disturbance during spawning, and positively related to dissolved soluble reactive phosphorus prior to spawning, which may indicate phosphorus limitation in the streams. These results suggest that enrichment from salmon nutrients does not always translate into elevated periphyton abundance. This underscores the need to directly assess the outcome of salmon impacts on streams rather than extrapolating from stable isotope evidence for the incorporation of salmon nutrients into food webs.     

Benthic algal biomass and productivity in high subarctic streams,Alaska

Year-round measurements of the standing crop of epilithic algae (as chlorophyll a concentration) in two streams — one second and one fourth order (map scale 1:63 360) — in interior Alaska (64°–65° N) were only about one tenth that reported from streams of temperate North America. Cell densities in these streams, however, were similar to those in comparable temperate streams. Year-round domination of the benthic flora by very tiny diatoms (Achnanthes spp.) may explain the apparent disparity between low chlorophyll a content and nearly average cell densities. Chlorophyll a standing crop in a more alkaline groundwater-fed stream, however, was higher and within the range of similarly sized temperate streams. Maximum chlorophyll a standing crop varied positively with alkalinity in 5 clear-water streams where standing crop was measured on natural or artificial substrates. Seasonal mean concentrations of sestonic chlorophyll a (used as estimates of benthic algal chlorophyll a standing crop) varied directly and significantly with alkalinity among ten clear-water streams; and, with total phosphorus among 8 of 10 clear-water and 5 brown-water streams studied. During the summer, when there is little darkness, gross primary productivity (as estimated by the diurnal dissolved-oxygen method) was similar to that of northern temperate streams. Gross primary productivity was also seen to vary directly with alkalinity in 5 clear-water streams of this region.U.S. Fish and Wildlife Service     

Applying the light : nutrient hypothesis to stream periphyton

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.     

Separating physical disturbance and nutrient enrichment caused by Pacific salmon in stream ecosystems

1. Pacific salmon (Oncorhynchus spp.) deliver marine‐derived nutrients to the streams in which they spawn and die, and these resource subsidies can increase the abundance of stream biota. In strong contrast, physical disturbance from salmon spawning activity can reduce the abundance of benthic organisms. Previous experimental designs have not established the relative effects of these two contrasting processes on stream organisms during a salmon run. 2. We combined manipulative and observational field studies to assess the degree of nutrient enrichment, physical disturbance, and the net effect of salmon on the abundance of benthic periphyton. Related salmon‐mediated processes were also evaluated for benthic macroinvertebrates. Mesh exclosures (2 × 2 m plots) prevented salmon from disturbing areas of the stream channel, which were compared with areas to which salmon had access. Sampling was conducted both before and during the late‐summer spawning run of pink (O. gorbushca) and chum (O. keta) salmon. 3. Streamwater nitrogen and phosphorus concentrations increased sharply with the onset of the salmon run, and highly significant positive relationships were observed between the numbers of salmon present in the stream and these dissolved nutrients. Before the salmon run, periphyton biomass (as chlorophyll a) and total macroinvertebrate abundance were very similar between control and exclosure plots. During the salmon run, exclosures departed substantially from controls, suggesting significant disturbance imparted on benthic biota. 4. Comparing exclosures before and during the salmon run enabled us to estimate the effects of salmon in the absence of direct salmon disturbance. This ‘nutrient enrichment potential’ was significant for periphyton biomass, as was a related index for macroinvertebrate abundance (although enhanced invertebrate drift into exclosures during the salmon run could also have been important). Interestingly, however, the net effect of salmon, evaluated by comparing control plots before and during the salmon run, was relatively modest for both periphyton and macroinvertebrates, suggesting that nutrient enrichment effects were largely offset by disturbance. 5. Our results illustrate the importance of isolating the specific mechanisms via which organisms affect ecosystems, and indicate that the relative magnitude of salmon nutrient enrichment and benthic disturbance determines the net effect that these ecologically important fish have on stream ecosystems.     

Impacts of an Invasive N2‐Fixing Tree on Hawaiian Stream Water Quality

N₂‐fixing trees can affect stream water quality. This has been documented in temperate streams, but not in tropical ones, even though N₂‐fixing trees are prevalent in the tropics. We investigated the effects of the introduced, invasive tree, Falcataria moluccana (albizia) on water quality of Hawaiian streams. Nutrient concentrations were measured in reaches above and below F. moluccana‐dominated riparian zones of four streams over 14 mo, and benthic algal nutrient limitation was examined in one stream. NO₃^?+NO₂^? concentrations were up to 600 percent higher in reaches below F. moluccana stands than in ones above them. In contrast, dissolved organic nitrogen concentrations were 24 percent lower in most reaches below F. moluccana stands, and NH₄⁺ and particulate nitrogen concentrations were similar above and below the stands. Dissolved organic carbon concentrations were up to 30 percent lower below F. moluccana stands, but particulate carbon concentrations were similar between reaches. Total dissolved phosphorus concentrations were similar above and below F. moluccana stands, whereas H₄SiO₄ concentrations were higher below the stands. In the stream where benthic chlorophyll a was measured, concentrations were three times higher below the F. moluccana stand than above it. Benthic algae were co‐limited by nitrogen and phosphorus above the F. moluccana stand, and limited by phosphorus below it. These results suggest that F. moluccana's presence relieved nitrogen‐limitation and caused the benthic algae to become solely phosphorus‐limited. Overall, our results demonstrate that F. moluccana can strongly affect the chemistry and primary producers of these tropical streams.     

Effects of dissolved organic matter and ultraviolet radiation on the accrual, stoichiometry and algal taxonomy of stream periphyton

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.     

Variation in responses to spawning Pacific salmon among three south-eastern Alaska streams

1. Pacific salmon are thought to stimulate the productivity of the fresh waters in which they spawn by fertilising them with marine‐derived nutrients (MDN). We compared the influence of salmon spawners on surface streamwater chemistry and benthic biota among three south‐eastern Alaska streams. Within each stream, reaches up‐ and downstream of barriers to salmon migration were sampled during or soon after spawners entered the streams. 2. Within streams, concentrations of dissolved ammonium and soluble reactive phosphorus (SRP), abundance of epilithon (chlorophyll a and ash‐free dry mass) and biomass of chironomids were significantly higher in reaches with salmon spawners. In contrast, biomass of the mayflies Epeorus spp. and Rhithrogena spp. was significantly higher in reaches lacking spawners. 3. Among streams, significant differences were found in concentrations of dissolved ammonium, dissolved organic carbon, nitrate and SRP, abundance of epilithon, and the biomass of chironomids and Rhithrogena. These differences did not appear to reflect differences among streams in spawner density, nor the changes in water chemistry resulting from salmon spawners. 4. Our results suggest that the ‘enrichment’ effect of salmon spawners (e.g. increased streamwater nutrient concentrations) was balanced by other concurrent effects of spawners on streams (e.g. sediment disturbance). Furthermore, the collective effect of spawners on lotic ecosystems is likely to be constrained by conditions unique to individual streams, such as temperature, background water chemistry and light attenuation.     

Limiting nutrients for periphyton growth in sub-alpine, forest, agricultural and urban streams

1. Nutrient-diffusing artificial substrata were used in summer and autumn to assess limiting nutrients for periphytic algal growth in streams draining sub-alpine, forested, agricultural and urban catchments in eastern Victoria, Australia. 2. Chlorophyll a density was primarily limited by nitrogen in most cases; often phosphorus was secondarily limiting. One sub-alpine, one forest and one agricultural stream were primarily phosphorus limited in at least one season. Added trace elements and vitamins did not increase chlorophyll density. 3. The dominant filamentous algal genera did not differ greatly between various nutrient enrichments. However, the relative abundance of Stigeodanium spp. was sometimes increased on substrata containing the limiting nutrient. 4. The results suggest that nutrient limitation is a widespread phenomenon in Victorian streams, and that limiting nutrients can be inferred from stream-water nitrogen to phosphorus ratios in many instances.     

How will increased temperature and nutrient enrichment affect primary producers in sub‐Arctic streams?

1. Spring‐fed streams, with temperatures ranging from 7.1 to 21.6 °C, in an alpine geothermal area in SW Iceland were chosen to test hypotheses on the effects of nutrients and temperature on stream primary producers. Ammonium nitrate was dripped into the lower reaches of eight streams, with higher reaches being used as controls, during the summers of 2006 and 2007. Dry mass of larger primary producers, epilithic chlorophyll a and biovolumes of epilithic algae were measured. 2. Bryophyte communities were dominated by Fontinalis antipyretica, and biomass was greatest in the warmest streams. Jungermannia exsertifolia, a liverwort, was found in low densities in few samples from cold streams but this species was absent from the warmest streams. 3. Nutrient enrichment increased the biomass of bryophytes significantly in warm streams. No effects of the nutrient addition were detected on vascular plants. The biomass of larger filamentous algae (mainly Cladophora spp.) was significantly increased by nutrient enrichment in cold streams but reduced by nutrients in warm streams. Thalloid cyanobacteria (Nostoc spp.) were not affected by nutrients in cold streams but decreased with nutrient addition in warm streams. Epilithic algal chlorophyll a was increased by nutrients in all streams and to a greater extent in 2007 than in 2006. Nutrient addition did not affect the epilithic chlorophyll a differently in streams of different temperatures. 4. There were small differential effects of nutrients, influenced by pH and conductivity, on different epilithic algal groups. 5. As global temperatures increase, animal husbandry and perhaps crop agriculture are likely to increase in Iceland. Temperature will directly influence the stream communities, but its secondary effects, manifested through agricultural eutrophication, are likely to be much greater.     

The contribution of flood disturbance, catchment geology and land use to the habitat template of periphyton in stream ecosystems

1. Periphyton chlorophyll a (chl a), ash-free dry mass, taxonomic composition, and cellular and water-column nutrients were analysed every 4 weeks for a year at sixteen stream sites in New Zealand. The hypothesis was investigated that broad-scale differences in mean monthly periphyton development are defined primarily by the frequency of flood disturbances and the periphyton's interaction with the nutrients. it us of the streams as determined by catchment geology and land use. 2. Overall, mean monthly chl a concentration declined with increasing flood frequency (r= -0.711, P < 0.001), and seasonality in chl a was better defined at sites with a low frequency of floods. Chlorophyll a concentration was generally low throughout the year at sites with frequent floods (> 15 yr^?1). 3. No relationship existed between inorganic nutrient concentrations and catchment geology or land development. However, conductivity declined significantly as a function of the percentage of the catchment underlain by nutrient-poor, hard rocks (plutonic and fine-grained metamorphic rocks) (r= -0.515, P < 0.05), but increased significantly with the percentage of the catchment in intensive agricultural land use (r= 0.799, P < 0.001). 4. Cellular nutrient concentrations suggested that nitrogen was the nutrient most commonly limiting periphyton production. In turn, cellular N concentrations declined significantly with increasing percentage of the catchment in hard rock (r= -0.5M, P < 0.05) and increased with percentage of the catchments in intensive agricultlural land use (r= 0.948, P < 0.001). 5.The sites were classified into three enrichment groups (high, moderate and low) based on their land use and underlying geology. Cellular N concentrations varied significantly among these enrichment groups (ANOVA F= 14.661, P < 0.001). 6. Log chl a decreased significantly with increases in the annual 80th percentile velocity. However, the relationship was significantly different among the enrichment groups. 7. A stepwise multiple regression on the full dataset identified that the frequency of floods, proportion of the catchment in high-intensity agricultural land use and proportion in alkaline rocks were the most significant factors explaining variation in mean monthly chl a among the sites (r²= 89%). 8. Overall, the results showed that flood disturbance and catchment enrichment regimes are probably the principal axes of the habitat template of periphyton among the study streams, and could be used to explain and predict broad-scale differences in periphyton development among other temperate stream ecosystems.     

Effects of nutrients and light on periphyton biomass and nitrogen uptake in Mediterranean streams with contrasting land uses

1. Nutrient diffusing substrata (NDS) were used to determine the relative importance of nutrients and light as potential limiting factors of periphyton biomass and nitrogen (N) uptake in Mediterranean streams subjected to different human impacts. The nutrients examined were phosphorus (P) and N, and we also further differentiated between the response of periphyton communities to N species (i.e. NO₃‐N and NH₄‐N). To examine the effect of light and nutrients on periphyton biomass, chlorophyll a accrual rates on NDS located at open and closed canopy sites were compared. The effect of nutrient availability on periphyton uptake was measured by ¹⁵N changes on the NDS after NO₃‐¹⁵N short‐term nutrient additions. 2. Results show that light was the main factor affecting algal biomass in the study streams. Algal biomass was in general higher at open than at closed canopy sites. Nutrient availability, as simulated with the NDS experiments, did not enhance algal biomass accrual in either of the 2 light conditions. 3. In the control treatments (i.e. ambient concentrations), periphyton NO₃‐N uptake rates increased and C : N molar ratios decreased consistently with increases in N availability across streams. NO₃‐N uptake rates were altered when ambient N concentrations were increased artificially in the N amended NDS. Periphyton assemblages growing on N enriched substrata seemed to preferentially take up N diffusing from the substratum rather than N from the water column. This response differed among streams, and depended on ambient N availability. 4. Periphyton biomass was not significantly different between substrata exposed to the two forms of available N sources. Nonetheless, we found differences in the effects of both N sources on the uptake of N from the water column. NH₄‐N seemed to be the preferred source of N for periphyton growing on NDS. 5. Results suggest that the effect of riparian zones on light availability, although seldom considered by water managers, may be more important than nutrients in controlling eutrophication effects derived from human activities. Finally, our results confirm that not only increases in concentration, but also stoichiometric imbalances should be considered when examining N retention in human altered streams.     

Relative importance of periphyton and phytoplankton in turbid and clear vegetated shallow lakes from the Pampa Plain (Argentina): a comparative experimental study

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.     

Ecological responses to nutrients in streams and rivers of the Colorado mountains and foothills

1. Abundance and composition of periphyton and benthic macroinvertebrates were treated as potential nutrient response variables for 74 streams in montane Colorado. The streams ranged from unenriched to mildly enriched with nutrients (N, P). 2. The study showed no meaningful relationship between periphyton biomass accumulation and concentrations of total or dissolved forms of nitrogen or phosphorus. Nutrient concentrations were also unrelated to periphyton and macroinvertebrate richness, diversity and community composition. Macroinvertebrate communities did, however, show a strong positive relationship to periphyton abundance. 3. A positive response of periphyton biomass to increasing nutrient concentrations has been well documented over large ranges of nutrient concentrations. Our study suggests that the nutrient response is suppressed by other controlling factors on the lower limb of the nutrient response curve (i.e. at low nutrient concentrations); a quantitatively significant response occurs only in excess of a threshold beyond which nutrients become dominant over other controlling factors. This interpretation of the results is consistent with published meta‐analyses showing lack of nutrient response for a high proportion of experimentally enriched periphyton communities, and division of responses between N and P for communities that do show growth in response to enrichment. 4. Grazing probably is not the key controlling variable for periphyton in Colorado mountain streams, given that the highest chlorophyll concentrations are associated with the highest abundances of macroinvertebrates. Modelling indicates that the initial amount of periphyton biomass at the start of the growing season, in conjunction with elevation‐related length of the growing season and water temperature, explains most of the variation in periphyton accumulation among these streams, but there is a yet unexplained suppression of periphyton growth rates across all elevations.     

A Note on Abiotic Factors that Constrain Periphyton Growth in Alpine Glacier Streams

Periphyton growth limitation experiments were conducted in five glacier streams during the main ice melt period in late summer using nutrient diffusing substrata (NDS) that contained nitrate and/or phosphate. Periphyton net growth was determined as chlorophyll a accrual after an exposure time of 4 weeks. In addition, primary water chemistry and physical parameters of the study streams were measured. These chemical and physical parameters characterised the sites as kryal (glacial) systems. Neither nutrient limitation nor a significant correlation between water chemistry and physical data and chlorophyll a values were apparent. A comparison between current velocity and ln‐transformed chlorophyll a values revealed a typical optimum curve with highest periphyton accrual at 0.5 m s^–1 on NDS. During the summer ablation period, the net growth of periphyton in these glacial streams appeared to be controlled primarily by current velocity.     

Effects of nutrient loading,temperature regime and grazing pressure on nutrient limitation of periphyton in experimental ponds

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The relative importance of shading and nutrients on algal production in subtropical streams

1. We used artificial substrata in forested and open streams in South-East Queensland, Australia, to determine the relative importance of shading from riparian vegetation and of nutrients on periphyton growth, and whether nitrogen and/or phosphorus limited algal productivity.
2. Nutrient-diffusing substrata consisting of agar enriched with N, P and N + P, and controls without nutrients, were deployed in duplicate at 15 sites in headwater streams with riparian canopy cover ranging from 0 to 88%.
3. Shading was the over-riding factor controlling periphyton biomass accrual on the artificial substrata, with nutrients playing a relatively minor role. Microscopic examination of periphyton scrapings taken after 7 weeks revealed that diatoms dominated on the artificial substrata in shaded streams, whereas filamentous green algae dominated the algal assemblage in the more open canopy streams.
4. Whilst nutrients had little effect on the accrual of algal biomass compared with riparian shading, there was evidence that nitrogen, and not phosphorus, stimulated periphyton production in streams with sufficient light.     

Responses of invertebrates and algae of a boreal coniferous forest stream to experimental manipulation of leaf litter inputs and shading

Forest harvesting alters leaf litter inputs and shading of small streams. Most of the previous studies of harvesting effects are limited to coastal or deciduous forests, so here we consider a sub-boreal forest stream. To test the hypothesis that changes in light and litter inputs would affect the benthic community in these streams, we experimentally manipulated these variables in stream mesocosms. We used a 2 × 2 factorial design with light (shaded or full light) and leaf litter inputs (equivalent to a forested stream or one quarter that rate) as factors. The high leaf litter treatment resulted in differences in macroinvertebrate community composition and higher densities of two shredders, Limnephilus sp. and Podmosta sp., suggesting food limitation. Algal filaments were longer in the high light treatments indicating a change in periphyton composition. There were no significant differences in chlorophyll a or ash-free dry mass, suggesting that light was not limiting to periphyton. The community structure clearly shifted in response to both resources, although primarily to detrital inputs. These results provide evidence that changes to shading and leaf inputs to small streams can affect the benthos and may limit secondary production.