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.     

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.     

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.     

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.     

Weedbeds and big bugs: the importance of scale in detecting the influence of nutrients and predation on macroinvertebrates in plant‐dominated shallow lakes

1. The scale of investigations influences the interpretation of results. Here, we investigate the influence of fish and nutrients on biotic communities in shallow lakes, using studies at two different scales: (i) within‐lake experimental manipulation and (ii) comparative, among‐lake relationships. 2. At both scales, fish predation had an overriding influence on macroinvertebrates; fish reduced macroinvertebrate biomass and altered community composition. Prey selection appeared to be size based. Fish influenced zooplankton abundance and light penetration through the water column also, but there was no indication that fish caused increased resuspension of sediment. 3. There were effects of nutrients at both scales, but these effects differed with the scale of the investigation. Nutrients increased phytoplankton and periphyton at the within‐lake scale, and were associated with increased periphyton at the among‐lake scale. No significant effect of nutrients on macroinvertebrates was observed at the within‐lake scale. However, at the among‐lake scale, nutrients positively influenced the biomass and density of macroinvertebrates, and ameliorated the effect of fish on macroinvertebrates. 4. Increased prey availability at higher nutrient concentrations would be expected to cause changes in the fish community. However, at the among‐lake scale, differences were not apparent in fish biomass among lakes with different nutrient conditions, suggesting that stochastic events influence the fish community in these small and relatively isolated shallow lakes. 5. The intensity of predation by fish significantly influences macroinvertebrate community structure of shallow lakes, but nutrients also play a role. The scale of investigation influences the ability to detect the influence of nutrients on the different components of shallow lake communities, particularly for longer lived organisms such as macroinvertebrates, where the response takes longer to manifest.     

Responses of lotic periphyton to pulses of phosphorus: P‐flux controlled growth rate

1. The response of stream periphyton to the addition of limiting nutrients has been the focus of many studies. However, the influence of pulsed nutrient additions has not previously been examined. This study investigates the biomass accrual and physiological responses of phosphorus‐limited lotic periphyton to hourly phosphate fluxes. 2. Two pulsing experiments were conducted: (i) a variable flux trial that compared variable hourly P‐fluxes, delivered either continuously at different concentrations or at the same concentration but in pulses of differing duration per hour and (ii) a constant flux trial that compared periphyton responses at a set hourly P‐flux but delivered in pulses of varying concentration and duration. 3. Growth response and alkaline phosphatase activity during the variable flux experiment showed that periphyton responds to the hourly integrated flux of phosphorus, regardless of whether the nutrient is supplied in short concentrated pulses or continuously at much lower concentrations. 4. The constant flux experiment examined the pulse period required to attain maximum biomass for a given phosphorus flux. Periphyton response to 5‐min pulses of phosphate per hour approximated the maximum biomass as that attained when the same hourly flux was added continuously. Compared with the control, there was also a substantial increase in biomass with pulses of only 1 min each hour. These results demonstrate that the hourly average phosphate concentration to which periphyton communities are exposed is paramount in determining P‐limited growth dynamics. 5. Species composition was not significantly different among treatments in each experiment; however, the design was to evaluate monotonic response with increasing phosphorus flux and species diversity may not respond monotonically. The data are therefore preliminary but suggest the need to determine if species diversity is generally lower when there are brief pulses of phosphate. Unlike pulse experiments that mimic lentic situations, nutrient additions were not used to completion and species success and composition was more dependent on their ability to acquire limiting nutrients rapidly rather than on their ability to take up nutrients at the lowest concentration.     

The role of light availability and herbivory on algal responses to nutrient enrichment in a riparian wetland,Alaska

We investigated how the relative availability of solar radiation in the presence or absence of grazing alters the ability of benthic algae to respond to nutrient enrichment in an Alaskan marsh. We used a factorial mesocosm experiment that included nutrient enrichment (enriched or control), grazing (grazed or ungrazed), and light (unshaded or shaded) to simulate shading by macrophytes early and late in the growing season, respectively. We found stronger effects of grazers and nutrients compared to light on benthic algal biomass and taxonomic composition. Algal biomass increased in nutrient‐enriched treatments and was reduced by grazing. Shading did not have an effect on algal biomass or taxonomic composition, but the concentration of chl a per algal biovolume increased with shading, demonstrating the ability of algae to compensate for changes in light availability. Algal taxonomic composition was more affected by grazer presence than nutrients or light. Grazer‐resistant taxa (basal filaments of Stigeoclonium) were replaced by diatoms (Nitzschia) and filamentous green algae (Ulothrix) when herbivores were removed. The interacting and opposing influences of nutrients and grazing indicate that the algal community is under dual control from the bottom‐up (nutrient limitation) and from the top‐down (consumption by herbivores), although grazers had a stronger influence on algal biomass and taxonomic composition than nutrient enrichment. Our results suggest that low light availability will not inhibit the algal response to elevated nutrient concentrations expected with ongoing climate change, but grazers rapidly consume algae following enrichment, masking the effects of elevated nutrients on algal production.     

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.     

Periphyton removal related to phosphorus and grazer biomass level

1. The proliferation of nuisance periphyton in enriched streams may be dependent on the biomass of the grazing macroinvertebrates present. In the present study, the effectiveness of grazer size and biomass in controlling periphyton and the extent to which grazing effectiveness was affected by enrichment level were determined. 2. Two sets of experiments with two caddisfly grazers were conducted in laboratory channels during spring and summer 1995 and 1996. The first set tested the combined effect of phosphorus enrichment and grazing, while the second set tested the effect of variable grazer biomass on periphyton biomass. 3. Grazing reduced periphyton biomass in excess of 80%, compared to ungrazed controls. Grazers were equally effective in controlling filamentous green algae, Stigeoclonium, diatoms and small colonial greens. Near complete removal of periphyton biomass by grazing occurred at even at the lowest grazer biomass level (750 mg m^??2, i.e. approximately one-third of natural levels). 4. Grazing controlled periphyton biomass more than did enrichment with soluble reactive phosphorus (SRP). 5. Grazing rates in the phosphorus-grazing interaction experiments averaged about 6 mg chl a g invertebrate^??1 day^??1, which was similar to past work in these channels and elsewhere, while rates were about five-fold higher in the variable grazer biomass experiments. 6. Simulating effects of SRP and grazing with a calibrated model suggests that higher SRP levels would be necessary to exceed a nuisance periphyton biomass level if grazers were present. However, if grazer biomass was more than 1500 mg m^??2, a nuisance level would probably not be exceeded at any SRP.     

Effects of nutrients and light on periphytic biomass and nutrient stoichiometry in a tropical black-water aquatic ecosystem

This article aims to test the light-nutrient hypothesis (LNH) in a periphytic community in a tropical black-water lake. Individual and interactive effects of light and nutrient availability were assessed with periphyton biomass accrual, nutrient content, and nutrient stoichiometry. We performed a manipulative field experiment with a 4 × 2 factorial design. We used nutrient diffusing substrates to produce four different nutrients treatments: Control (no nutrient added), nitrogen amended (N), phosphorus amended (P) and combined N and P amendment (NP). Two light levels were also considered: high light (near surface water) and low light (near bottom water). Light and nutrients individually and interactively caused significant changes in aggregate periphyton community properties. Total and autotrophic biomasses were significantly higher in high light conditions and in nutrient enriched treatments. Autotrophic biomass was significantly higher in N enriched treatment whereas total biomass was mainly affected by the joint addition of N and P. At lower light availability periphyton growth was limited, even in enriched treatments. Light also strongly affected periphyton nutrient content. Periphyton C, N and P in general increased when subjected to high light conditions. As predicted by the LNH, light promoted an increase in periphyton C:P ratios in P deprived treatments, but an opposite effect was observed on C:N ratios, especially in N-enriched treatments. This experiment revealed that light availability strongly limits the propagation of nutrient effects on periphyton growth. Such complex interdependencies on basal resources affect the proportion of autotrophic to total periphytic biomass that can be an important mechanism to explain variation in the nutrient stoichiometry of periphyton in nature.     

Spatial variation in periphyton fatty acid composition in subtropical streams

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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.     

Response of Periphytic Algae to Gradients in Nitrogen and Phosphorus in Streamside Mesocosms

In this study we manipulated both nitrogen and phosphorus concentrations in stream mesocosms to develop quantitative relationships between periphytic algal growth rates and peak biomass with inorganic N and P concentrations. Stream water from Harts Run, a 2nd order stream in a pristine catchment, was constantly added to 36 stream-side stream mesocosms in low volumes and then recirculated to reduce nutrient concentrations. Clay tiles were colonized with periphyton in the mesocosms. Nutrients were added to create P and N concentrations ranging from less than Harts Run concentrations to 128 μg SRP l⁻¹ and 1024 μg NO₃-N l⁻¹. Algae and water were sampled every 3 days during colonization until periphyton communities reached peak biomass and then sloughed. Nutrient depletion was substantial in the mesocosms. Algae accumulated in all streams, even streams in which no nutrients were added. Nutrient limitation of algal growth and peak biomass accrual was observed in both low P and low N conditions. The Monod model best explained relationships between P and N concentrations and algal growth and peak biomass. Algal growth was 90% of maximum rates or higher in nutrient concentrations 16 μg SRP l⁻¹ and 86 μg DIN l⁻¹. These saturating concentrations for growth rates were 3–5 times lower than concentrations needed to produce maximum biomass. Modified Monod models using both DIN and SRP were developed to explain algal growth rates and peak biomass, which respectively explained 44 and 70% of the variance in algal response.     

The influence of dissolved nutrients and particulate organic matter quality on microbial respiration and biomass in a forest stream

1. Although dissolved nutrients and the quality of particulate organic matter (POM) influence microbial processes in aquatic systems, these factors have rarely been considered simultaneously. We manipulated dissolved nutrient concentrations and POM type in three contiguous reaches (reference, nitrogen, nitrogen + phosphorus) of a low nutrient, third‐order stream at Hubbard Brook Experimental Forest (U.S.A). In each reach we placed species of leaves (mean C : N of 68 and C : P of 2284) and wood (mean C : N of 721 and C : P of 60 654) that differed in elemental composition. We measured the respiration and biomass of microbes associated with this POM before and after nutrient addition. 2. Before nutrient addition, microbial respiration rates and biomass were higher for leaves than for wood. Respiration rates of microbes associated with wood showed a larger response to increased dissolved nutrient concentrations than respiration rates of microbes associated with leaves, suggesting that the response of microbes to increased dissolved nutrients was influenced by the quality of their substrate. 3. Overall, dissolved nutrients had strong positive effects on microbial respiration and fungal, but not bacterial, biomass, indicating that microbial respiration and fungi were nutrient limited. The concentration of nitrate in the enriched reaches was within the range of natural variation in forest streams, suggesting that natural variation in nitrate among forest streams influences carbon mineralisation and fungal biomass.     

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.     

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.     

The effect of different nutrient concentrations on the growth rate and nitrogen storage of watercress (Nasturtium officinale R. Br.)

The mechanisms that allow broadly distributed aquatic plants to inhabit variable resource environments are unclear, yet understanding these mechanisms is important because broad environmental tolerance is often linked to invasiveness in terrestrial and aquatic plants. In an experimental stream, we examined the effects of different nutrient concentrations on the growth rate, biomass, and foliar nutrient concentrations of a cosmopolitan and potentially invasive aquatic plant, Nasturtium officinale (R. Br.). Nasturtium seedlings were grown under six nutrient treatment levels ranging from 0.64 μm N:0.09 μm P to 1531 μm N:204.13 μm P, for 8 weeks. Absolute and relative growth rates, and biomass of seedlings increased along a gradient of increasing nutrient concentrations but the effect of nutrient concentration was dependent on growing time. Seedling biomass varied among nutrient treatments in weeks 4 through 8 of the experiment, but did not differ in week 2. By week 8, the two highest nutrient treatments had greater biomass than the two lowest nutrient treatments. Foliar nitrogen concentration increased, whereas carbon concentration and C:N ratios decreased in response to increasing nutrients. Nasturtium grows slowly in nutrient-poor conditions but rapidly increases its growth, biomass accrual, and nitrogen storage as conditions become nutrient-rich. The response of Nasturtium to enhanced nutrient conditions may indicate how aquatic nuisance species successfully invade and dominate plant communities in streams, where resources often vary both temporally and spatially.     

Responses of stream algae to grazing minnows and nutrients: a field test for interactions

Summary Previous studies have shown that an algivorous grazing minnow (Campostoma anomalum) is the major herbivore in Brier Creek, a hardwater stream in south central Oklahoma. In summer and autumn schools of Campostoma virtually eliminate algae from substrate surfaces in deeper areas of some pools. The pool-to-pool distributions of algae and Campostoma reported for this stream could occur if nutrient limitation permits grazing by Campostoma to outrun algal growth. To test this hypothesis, mesh pens were built to exclude Campostoma from substrates in each of four typical Campostoma pools. N+P+K lawn fertilizer was added daily to two of the four pools; the other two, which received no fertilizer additions and which were not visibly affected by fertilizer transported downstream from the pools enriched with nutrients, served as controls. Algae accumulated rapidly on natural substrates and on unglazed ceramic tiles in grazer-exclusion pens in pools receiving N+P+K additions and more slowly in pens in both control pools. Periphyton biomass on grazed substrates in all four pools remained low throughout the experiment. Hence, Campostoma at normal densities were able to outrun algal growth even when nutrients were added. Eleven days after the experiment started, I determined biomass, biomass-specific net primary productivity, and areal net primary productivity of periphyton on substrates exposed to all combinations of grazer (+,0) and nutrient (+,0) treatments. Grazing increased biomass-specific primary productivity, prevented accumulation of biomass, and decreased areal primary productivity of periphyton. Additions of N+P+K increased biomass-specific net primary productivity of grazed and ungrazed periphyton and markedly increased biomass of periphyton on substrates protected from Campostoma. Although food supply for Campostoma appeared to be greater with nutrient additions, condition of Campostoma in pools receiving N+P+K was not significantly different from Campostoma collected from control pools 35 days after the experiment started. I conclude that although nutrient supply limits biomass-specific primary productivity of periphyton in Brier Creek, nutrient limitation in this stream exacerbates, rather than causes, the visually conspicuous pool-to-pool complimentary distribution of algae and Campostoma: in this stream, grazing by Campostoma at natural densities can outrun periphyton growth even when nutrients are added.     

Non-native earthworms in riparian soils increase nitrogen flux into adjacent aquatic ecosystems

Riparian zones are an important transition between terrestrial and aquatic ecosystems, and they function in nutrient cycling and removal. Non-native earthworms invading earthworm-free areas of North America can affect nutrient cycling in upland soils and have the potential to affect it in riparian soils. We examined how the presence of earthworms can affect riparian nutrient cycling and nutrient delivery to streams. Two mesocosm experiments were conducted to determine how (1) the biomass of earthworms and (2) earthworm species can affect nutrient flux from riparian zones to nearby streams and how this flux can affect streamwater nutrients and periphyton growth. In separate experiments, riparian soil cores were amended with one of four mixed earthworm biomasses (0, 4, 10, or 23 g m(-2) ash-free dry mass) or with one of three earthworm species (Aporrectodea caliginosa, Lumbricus terrestris, L. rubellus) or no earthworm species. Riparian soil cores were coupled to artificial streams, and over a 36-day period, we measured nutrient leaching rates, in-stream nutrient concentrations, and periphyton growth. Ammonium leaching increased with increasing biomass and was greatest from the A. caliginosa treatments. Nitrate leaching increased through time and increased at a greater rate with higher biomass and from cores containing A. caliginosa. We suggest that the overall response of increased nitrate leaching [90% of total nitrogen (N)] was due to a combination of ammonium excretion and burrowing by earthworms, which increased nitrification rates. During both experiments, periphyton biomass increased through time but did not differ across treatments despite high in-stream inorganic N. Through time, in-stream phosphorus (P) concentration declined to <5 microg l(-1), and periphyton growth was likely P-limited. We conclude that activities of non-native earthworms (particularly A. caliginosa) can alter biogeochemical cycling in riparian zones, potentially reducing the N-buffering capacity of riparian zones and altering stoichiometric relationships in adjacent aquatic ecosystems.     

Antarctic Stream Ecosystems: Variability in Environmental Properties and Algal Community Structure

The variability in physical, chemical and biological properties was examined for a number of glacier melt streams in south Victoria Land, Antarctica. Streams flowed for between one and two months. Stream water temperatures (range = 0–11°C) varied over short (hr) time scales whilst discharges varied considerably between streams (range 0.001–15 m³s⁻¹) and over diel cycles. Solar radiation and air temperature were major determinants of stream discharge. Variability in discharge was reflected in variability in nutrient chemistry and sediment load. Nitrogen and phosphorus varied considerably between streams; the meltwaters early in summer contained 10–20 times higher levels of dissolved N and P than later in the season. Within stream nutrient levels were modified by dense algal growths and penguin rookeries. Epilithic algal communities were made up predominantly of cyanophyceae which formed mats and crusts. Longitudinal and horizontal variability of species in the communities in selected streams is described. Analyses of algal cover and biomass (chlorophyll a) show that substrate type and flow rates are of greater importance than nutrients in influencing algal abundance and biomass. In some streams biomass values of over 20 μg Ch. a cm⁻² were recorded, much of which remains viable but inactive over the antarctic winter.