Seasonal shifts in the importance of bottom–up and top–down factors on stream periphyton community structure

We examined the importance of temporal variability in top–down and bottom–up effects on the accumulation of stream periphyton, which are complex associations of autotrophic and heterotrophic microorganisms. Periphyton contributes to primary production and nutrient cycling and serves as a food resource for herbivores (grazers). Periphyton growth is often limited by the availability of nitrogen and phosphorus, and biomass can be controlled by grazers. In this study we experimentally manipulated nutrients and grazers simultaneously to determine the relative contribution of bottom–up and top–down controls on periphyton over time. We used nutrient diffusing substrates to regulate nutrient concentrations and an underwater electric field to exclude grazing insects in three sequential 16–17 day experiments from August to October in montane Colorado, USA. We measured algal biomass, periphyton organic mass, and algal community composition in each experiment and determined densities of streambed insect species, including grazers. Phosphorus was the primary limiting nutrient for algal biomass, but it did not influence periphyton organic mass across all experiments. Effects of nutrient additions on algal biomass and community composition decreased between August and October. Grazed substrates supported reduced periphyton biomass only in the first experiment, corresponding to high benthic abundances of a dominant mayfly grazer (Rhithrogena spp.). Grazed substrates in the first experiment also showed altered algal community composition with reduced diatom relative abundances, presumably in response to selective grazing. We showed that top–down grazing effects were strongest in late summer when grazers were abundant. The effects of phosphorus additions on algal biomass likely decreased over time because temperature became more limiting to growth than nutrients, and because reduced current velocity decreased nutrient uptake rates. These results suggest that investigators should proceed with caution when extending findings based on short‐term experiments. Furthermore, these results support the need for additional seasonal‐scale field research in stream ecology.     

Elevated light and nutrients alter the nutritional quality of stream periphyton

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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.     

High nutrient availability leads to weaker top‐down control of stream periphyton: Compensatory feeding in Ancylus fluviatilis

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Nutrient loading and grazing by the minnow Phoxinus erythrogaster shift periphyton abundance and stoichiometry in mesocosms

1. Anthropogenic activities in prairie streams are increasing nutrient inputs and altering stream communities. Understanding the role of large consumers such as fish in regulating periphyton structure and nutritional content is necessary to predict how changing diversity will interact with nutrient enrichment to regulate stream nutrient processing and retention. 2. We characterised the importance of grazing fish on stream nutrient storage and cycling following a simulated flood under different nutrient regimes by crossing six nutrient concentrations with six densities of a grazing minnow (southern redbelly dace, Phoxinus erythrogaster) in large outdoor mesocosms. We measured the biomass and stoichiometry of overstory and understory periphyton layers, the stoichiometry of fish tissue and excretion, and compared fish diet composition with available algal assemblages in pools and riffles to evaluate whether fish were selectively foraging within or among habitats. 3. Model selection indicated nutrient loading and fish density were important to algal composition and periphyton carbon (C): nitrogen (N). Nutrient loading increased algal biomass, favoured diatom growth over green algae and decreased periphyton C : N. Increasing grazer density did not affect biomass and reduced the C : N of overstory, but not understory periphyton. Algal composition of dace diet was correlated with available algae, but there were proportionately more diatoms present in dace guts. We found no correlation between fish egestion/excretion nutrient ratios and nutrient loading or fish density despite varying N content of periphyton. 4. Large grazers and nutrient availability can have a spatially distinct influence at a microhabitat scale on the nutrient status of primary producers in streams.     

The influence of crevice size on the protection of epilithic algae from grazers

1. This study investigated how the size of crevices might affect their effectiveness as refuges for diatom‐dominated algal assemblages from the grazing minnow Campostoma anomalum. 2. Crevice size was manipulated by making fired‐clay substrates, using moulds to produce eight substrates with pits from 1.17 to 22.0 mm diameter. Non‐pitted clay‐stones and limestone were also tested. Cages were used to control the access of Campostoma to arrays of the 10 different algal‐colonised substrates. The grazing treatments were: open and grazed, caged and ungrazed, and a grazed cage control. The experiment was replicated in eight large outdoor tanks. After 3 weeks, substrates were brushed and chlorophyll a concentrations of the removed algae and the algae remaining in pits were measured. 3. The experiment was field‐validated by exposing arrays of substrates to grazing Campostoma in five pools of a limestone stream. 4. The clay‐stone and limestone substrates accrued similar algal biomass and assemblages. 5. Smaller crevices provided more protection against grazing than larger crevices. Specifically, pits with openings smaller than 2 mm protected the enclosed algal assemblages in both the tank and field experiments. Larger pits provided less protection and pits over 7 mm in diameter were heavily grazed and may even be preferentially grazed by Campostoma. 6. None of the tested pit sizes were protective against larval chironomid grazers in the tank experiment, demonstrating that differences in the grazer size influence the effectiveness of crevice refuges.     

Light- and nutrient-limited periphyton in low order streams of Oahu,Hawaii

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

Periphytic food and predatory crayfish: relative roles in determining snail distribution

Summary In the laboratory and field, we examined how periphyton (food of snails) and predatory crayfish influenced snail distribution in Trout Lake, a permanent, northern Wisconsin lake. Laboratory experiments (with no crayfish) tested the importance of periphyton biomass in determining snail preference among rocks, and among rock, sand, and macrophyte substrates. Among rocks with four different amounts of periphyton, periphyton biomass and the number of Lymnaea emarginata, Physa spp., and Amnicola spp. were positively related. A similar, but non-significant, trend occurred for Helisoma anceps. A field experiment at a site in Trout Lake where predation risk was low confirmed the preference by snails for periphyton covered rocks; more snails colonized rocks with periphyton than rocks without. When given a choice of rock, sand, and macrophytes in the laboratory, L. emarginata preferred high periphyton biomass and rock. Laboratory and field results contrasted with the distribution of snails in Trout Lake; no snails occurred in areas with abundant periphyton-covered rocks, but snails were abundant nearby on scattered rocks with little periphyton. However, where snails were absent, crayfish were abundant (14.5 crayfish-trap^–1-day^–1), and where snails were abundant, crayfish were rare (3.2 crayfish-trap^–1-day^–1), suggesting that crayfish predation reduced snails. The hypothesis that the negative association between snail and periphyton biomass resulted from snail grazing was supported by the results of a field snail enclosure-exclosure experiment (1 m² cages; n=3). All experiments and observations therefore suggest that: 1) crayfish predation is more important than a preference for high periphyton biomass in determining snail distribution in Trout Lake; 2) periphyton biomass is negtively related to snail grazing; and 3) crayfish had a positive indirect effect on periphyton by preying on grazing snails.     

Feedbacks of consumer nutrient recycling on producer biomass and stoichiometry: separating direct and indirect effects

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

Effects of copper on periphyton communities assessed in situ using chemical-diffusing substrates

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

Depth distribution of Campostoma grazing scars in an Ozark stream

Synopsis Campostoma spp., widespread and abundant herbivorous minnows of eastern North America, produce distinctive grazing scars when feeding on algae attached to natural substrates in streams. These scars are particularly prominent upon the low growth forms of blue-green algae that dominate the attached algal flora of many upland streams. In one stream pool in the Ozark uplands of Oklahoma, numbers and sizes of grazing scars coincided with numbers and sizes of individual Campostoma that occurred across a depth gradient, demonstrating that the information contained in the scars can provide quantification of microhabitat use and grazing intensity of these important herbivores. The results also support the hypothesis that in environments free of aquatic predators, larger fish use deeper parts of available stream habitats, particularly if threats from terrestrial or avian predators exist.     

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

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Nutrient Limitation to Primary Productivity in a Secondary Savanna in Venezuela1

We examined nutrient limitation to primary productivity in a secondary savanna in the interior branch of the Coastal Range of Venezuela, which was converted from forest to savanna more than 100 years ago. We manipulated soil nutrients by adding nitrogen (+N), phosphorus and potassium (+PK), and nitrogen, phosphorus, and potassium (+NPK) to intact savanna. Eleven months after fertilization, we measured aboveground biomass and belowground biomass as live fine roots in the top 20 cm of soil, and species and functional group composition in response to nutrient additions. Aboveground biomass was highest in the NPK treatment ([mean g/m2]; control = 402, +N = 718, +PK = 490, +NPK = 949). Aboveground production, however, appeared to be limited primarily by N. Aboveground biomass increased 78 percent when N was added alone but did not significantly respond to PK additions when compared to controls. In contrast to aboveground biomass, belowground biomass increased with PK additions but showed no significant increase with N (depth 0–20 cm; [mean g/m2]; control = 685, +N = 443, +PK = 827, +NPK = 832). There was also a 36 percent increase in root length with PK additions when compared to controls. Whole savanna shoot:root ratios were similar for control and +PK (0.6), while those for +N or +NPK fertilization were significantly higher (1.7 and 1.2, respectively). Total biomass response (above + belowground) to nutrient additions showed a strong N and PK co‐limitation ([mean g/m2]; control = 1073, +N = 1111, +PK = 1258, +NPK = 1713). Aboveground biomass of all monocots increased with N additions, whereas dicots showed no response to nutrient additions. Trachypogon spp. (T. plumosus+T. vestitus) and Axonopus canescens, the two dominant grasses, made up more than 89 percent of the total aboveground biomass in these sites. Trachypogon spp. responded to NPK, whereas A. canescens, sedges, and the remaining monocots only responded to N. Even though nutrient additions resulted in higher aboveground biomass in N and NPK fertilized plots, this had little effect on plant community composition. With the exception of sedges, which responded positively to N additions and increased from 4 to 8 percent of die plant community, no changes were observed in plant community composition after 11 months.     

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 23 μg · L^?1 and significantly decreased with increasing SO₄. Mean chl a concentrations at sites ranged from 0.2 to 8.1 μg · 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 on the 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.     

Grazing and the distribution of sediment particle sizes in artificial stream systems

Recent research has shown that grazing by the algivorous minnows of the genus Campostoma can have a significant effect on both structural and functional parameters in stream ecosystems, influencing algal height and type, primary productivity, carbon dynamics, bacterial biomass and the size fractionation of benthic organic matter. This study was undertaken to evaluate the effect of minnow grazing on benthic particulate organic matter (BPOM) under the controlled conditions available in experimental streams. For this study, four grazed and four ungrazed artificial streams were sampled for BPOM on two dates. The samples were partitioned into large, medium, fine and ultrafine fractions via wet filtration. Fish grazed systems had significantly larger percentages of the fine fraction and significantly smaller percentages of the ultrafine fraction. The ability of grazers to alter particle size distribution is important to overall stream organic matter dynamics because recent studies have shown the importance of particle size in determining bacterial numbers and activity, which in turn influence fundamental stream process like respiration and organic carbon dynamics.     

Factors influencing biomass and nutrient content of the submersed macrophyte Egeria densa Planch. in a pampasic stream

We identified factors influencing biomass and nutrient content in E. densa in an enriched pampean stream of Argentina. Physical (current velocity, temperature), chemical (pH, conductivity, dissolved oxygen, nutrient content in water and sediments), and biological variables (biomass and nutrient content of E. densa, biomass of periphyton and other macrophytes) were estimated at each sampling occasion, and mean monthly values estimated. Biomass and nutrient content in E. densa were correlated with these physical-chemical and biological variables. Biomass was positively correlated with ammonium in stream water (P<0.05) and sediment total nitrogen (P<0.01). Nitrogen showed a positive relationship with ammonium (P<0.01), and a negative one with nitrate and periphyton biomass (P<0.05). Phosphorus was positively correlated with soluble reactive phosphorus (P<0.01). The growth of other macrophyte species in the stream seemed to influence E. densa biomass, probably through competition for light. Current velocity was low and not significantly related with E. densa biomass, however, a flood at the beginning of the study washed the macrophyte stand downstream.     

Effect of periphyton biomass on hydraulic characteristics and nutrient cycling in streams

The effect of periphyton biomass on hydraulic characteristics and nutrient cycling was studied in laboratory streams with and without snail herbivores. Hydraulic characteristics, such as average water velocity, dispersion coefficients, and relative volume of transient storage zones (zones of stationary water), were quantified by performing short-term injections of a conservative tracer and fitting an advection-dispersion model to the conservative tracer concentration profile downstream from the injection site. Nutrient cycling was quantified by measuring two indices: (1) uptake rate of phosphorus from stream water normalized to gross primary production (GPP), a surrogate measure of total P demand, and (2) turnover rate of phosphorus in the periphyton matrix. These measures indicate the importance of internal cycling (within the periphyton matrix) in meeting the P demands of periphyton. Dense growths of filamentous diatoms and blue-green algae accumulated in the streams with no snails (high-biomass streams), whereas the periphyton communities in streams with snails consisted almost entirely of a thin layer of basal cells of Stigeoclonium sp. (low-biomass streams). Dispersion coefficients were significantly greater and transient storage zones were significantly larger in the high-biomass streams compared to the low-biomass streams. Rates of GPP-normalized P uptake from water and rates of P turnover in periphyton were significantly lower in high biomass than in low biomass periphyton communities, suggesting that a greater fraction of the P demand was met by recycling in the high biomass communities. Increases in streamwater P concentration significantly increased GPP-normalized P uptake in high biomass communities, suggesting diffusion limitation of nutrient transfer from stream water to algal cells in these communities. Our results demonstrate that accumulations of periphyton biomass can alter the hydraulic characteristics of streams, particularly by increasing transient storage zones, and can increase internal nutrient cycling. They suggest a close coupling of hydraulic characteristics and nutrient cycling processes in stream ecosystems.     

Effects of irradiance on diel and seasonal patterns of nutrient uptake by stream periphyton

1. Irradiance strongly affects the abundance of stream periphyton communities that in turn influence patterns of instream nutrient uptake. We examined relationships between irradiance and periphyton nutrient uptake taking into account diel and seasonal variation in ambient irradiance. 2. Uptake of dissolved N, P and C by periphyton as areal uptake (U) and demand (V_f) was determined under 11 irradiance levels (0–100% of ambient conditions) using shallow stream‐side experimental channels. Experiments were conducted once per season over one annual cycle with both day and night uptake rates assessed, together with periphyton biomass and autotrophic production rates. 3. No consistent diel variation in areal uptake or demand was detected for the predominant inorganic or total dissolved nutrients even at the highest irradiances. Lack of variation may indicate nutrient limitation, with photosynthetic sequestration and storage of C during the day for subsequent utilisation at night. Alternatively, oxygen consumption by photoautotrophs at night may stimulate compensatory heterotrophic uptake (e.g. denitrification). 4. In all seasons, release of dissolved organic N was detected during the day but to a lesser extent at night. This was not directly related to irradiance levels, indicating that heterotrophic metabolism (e.g. microbial decomposition) contributes to this phenomenon. 5. Areal uptake and demand for the predominant inorganic and total dissolved nutrients increased in response to increasing irradiance in some or all seasons, but rates were typically higher during the spring and summer. Saturation of areal uptake and demand at elevated irradiances was evident during the spring. demand was also saturated at higher irradiances in the summer and autumn. Maximum demand was comparable during spring and summer, but saturation occurred at lower irradiance in summer (24 h average 135–145 μmol m^?2 s^?1) relative to spring (312–424 μmol m^?2 s^?1), indicating more efficient nutrient uptake in summer. Higher total periphyton biomass in summer, but comparable autotrophic biomass (chlorophyll a), implies that heterotrophic metabolism may contribute to this greater efficiency. In spring, autotrophic biomass peaked at an irradiance level of 225 μmol m^?2 s^?1, also suggesting a role for heterotrophic metabolism in demand at higher irradiances. 6. The results of this study show that irradiance levels exert a strong influence on the nature and quantity of instream nutrient uptake with N demand saturated at elevated irradiance levels during the spring, summer and autumn. Our results also suggest that heterotrophic metabolism makes a measurable contribution to instream nutrient uptake even under higher irradiances that favour autotrophic activity.     

Habitat-Specific Nitrogen Dynamics in New Zealand Streams Containing Native or Invasive Fish

Streams are important sites of nutrient transport and transformation in the landscape but little is known about the way in which individual taxa or individual habitats (riffles and pools) influence nutrient dynamics within stream reaches. We used 5-week additions of a stable isotope (¹⁵NH₄Cl) tracer to investigate nitrogen dynamics in pools and riffles of two New Zealand streams, one with native fish (Galaxias depressiceps) and the other with invasive brown trout (Salmo trutta). In New Zealand, brown trout initiate a trophic cascade leading to increased algal biomass that we predicted would lead to higher N uptake and retention. Uptake of NO₃, but not ammonium, was greater in the trout stream. Rather than causing a large increase in N demand, trout may induce a reallocation of N uptake and retention among food web compartments in different habitats. The largest differences between streams were apparent in riffles, where most uptake and retention of N occurred. In the trout stream, uptake rate by epilithon in riffles was more than six times greater than uptake rates of any other compartment. In the Galaxias stream, several compartments in both habitats had similar uptake rates. Epilithon also accounted for a larger percentage of the ¹⁵N retained in the study reach in the trout stream (51%) than the Galaxias stream (34%). Our results show that an individual predatory taxon (in our case an invader) can influence N dynamics in streams but that the magnitude and location of the impact depend on a range of abiotic and biotic factors involved in N dynamics in streams.     

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

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