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.     

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.     

Fluctuations des nutriments au cours de la marée sur les parcs Ostreicoles de la rivière Penze (Nord-Finistère)

The design and performance of a simple, community level ecotoxicological testsystem is reported. Samples of periphyton communities, established on artificial substratum in natural streams were used to study effects on photosynthetic activity in short-term experiments. Photosynthesis was measured as light-dependent oxygen evolution or as ¹⁴CO₂-incorporation. The variability in photosynthetic activity between samples collected at the same time, expressed as coefficient of variation, was ca 20%. The variation in sensitivity of periphyton photosynthesis as dependent on sampling season was less than threefold for the two long-chained aliphatic amines and the textile industry effluent studied. Effects of the amines on periphyton from five different streams were also investigated. The ratio between maximum and minimum values of sensitivity was 5.6. It is concluded that the variation in sensitivity between different periphyton communities is similar to or less than that observed for fresh-water algal species. Some advantages with regard to ecological realism of using periphyton communities as test systems are discussed.     

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.     

Influence of suspended clay on phosphorus uptake by periphyton

We investigated the effect of suspended clay upon the phosphorus uptake rate exhibited by lotic periphyton communities. Suspended inorganic clays and periphyton are common to aquatic environments, and both can strongly influence physical and chemical water conditions. We used replicated artificial stream channels to test the prediction that suspended clay particles would affect the uptake of soluble reactive phosphorus (SRP) by periphyton. Commercially available kaolinite and bentonite clays were characterized for their aqueous suspension behavior and affinities for SRP. Periphyton was grown in a recirculating stream system and subjected to simultaneous suspended clay and SRP additions. SRP removal from solution, both in the presence and absence of suspended clays, was used to quantify SRP uptake parameters by periphyton. Clay type and concentrations of 20, 80, and 200 mg l⁻¹ had no significant effect upon SRP uptake rate exhibited by periphyton during three 90-min experiments. Less than 1% of SRP removal was attributable to the suspended clay load or artificial stream construction materials, based on clay isotherm data and material sorption studies, indicating that 99% of SRP removal was attributable to biotic uptake. Removal of SRP (as KH₂PO₄) was described by a first-order equation with rate constants ranging between 0.02 and 0.14 min⁻¹. Our results suggest that high turbidity conditions caused by suspended mineral clays have little immediate effect upon SRP removal from the water column by periphyton. Handling editor: D. Ryder     

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.     

Phosphate limitation influences the sensitivity to copper in periphytic algae

1. To assess the influence of nutrient limitation on copper toxicity, periphitic communities from an oligotrophic stream were exposed to copper for six to 12 days with and without the supply of nutrients (mainly P). 2. In contrast to the hypothesis that nutrient cycling in mature biofilms would protect them from Cu toxicity, low and high biomass biofilms did not differ in their physiological response to copper after 6 days of exposure. 3. A clear influence of P‐limitation on copper toxicity was observed. Periphytic communities that were previously fertilised for 18 days were three times more tolerant than control communities indicating that P‐limitation enhanced Cu toxicity and tolerance induction were probably related to the higher P‐availability. In addition, a compensation of Cu toxicity after P‐addition was observed in the long‐term (after 12 days). 4. We conclude that periphyton from oligotrophic streams is more sensitive to copper than periphyton from fertilised streams, and that therefore a higher effect of chronic copper exposure is expected to occur in oligotrophic P‐limited fluvial systems. 5. Extrapolation of our results to the area of study (Catalonia, north‐east of Spain) indicates that while the levels of Cu commonly found in the zone may negatively affect the periphyton from oligotrophic streams, because of the interaction between Cu and P, they are not able to control the growth of nuisance algae which is common under high nutrient conditions.     

Predator-Driven Nutrient Recycling in California Stream Ecosystems

Nutrient recycling by consumers in streams can influence ecosystem nutrient availability and the assemblage and growth of photoautotrophs. Stream fishes can play a large role in nutrient recycling, but contributions by other vertebrates to overall recycling rates remain poorly studied. In tributaries of the Pacific Northwest, coastal giant salamanders (Dicamptodon tenebrosus) occur at high densities alongside steelhead trout (Oncorhynchus mykiss) and are top aquatic predators. We surveyed the density and body size distributions of D. tenebrosus and O. mykiss in a California tributary stream, combined with a field study to determine mass-specific excretion rates of ammonium (N) and total dissolved phosphorus (P) for D. tenebrosus. We estimated O. mykiss excretion rates (N, P) by bioenergetics using field-collected data on the nutrient composition of O. mykiss diets from the same system. Despite lower abundance, D. tenebrosus biomass was 2.5 times higher than O. mykiss. Mass-specific excretion summed over 170 m of stream revealed that O. mykiss recycle 1.7 times more N, and 1.2 times more P than D. tenebrosus, and had a higher N:P ratio (8.7) than that of D. tenebrosus (6.0), or the two species combined (7.5). Through simulated trade-offs in biomass, we estimate that shifts from salamander biomass toward fish biomass have the potential to ease nutrient limitation in forested tributary streams. These results suggest that natural and anthropogenic heterogeneity in the relative abundance of these vertebrates and variation in the uptake rates across river networks can affect broad-scale patterns of nutrient limitation.     

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.     

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.     

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.     

Catabolic diversity of periphyton and detritus microbial communities in a subtropical wetland

The catabolic diversity of wetland microbial communities may be a sensitive indicator of nutrient loading or changes in environmental conditions. The objectives of this study were to assess the response of periphyton and microbial communities in water conservation area-2a (WCA-2a) of the Everglades to additions of C-substrates and inorganic nutrients. Carbon dioxide and CH₄ production rates were measured using 14 days incubation for periphyton, which typifies oligotrophic areas, and detritus, which is prevalent at P-impacted areas of WCA-2a. The wetland was characterized by decreasing P levels from peripheral to interior, oligotrophic areas. Microbial biomass and N mineralization rates were higher for oligotrophic periphyton than detritus. Methane production rates were also higher for unamended periphyton (80 mg CH₄-C kg⁻¹ d⁻¹) than detritus (22 mg CH₄-C kg⁻¹ d⁻¹), even though the organic matter content was higher for detritus (80%) than periphyton (69%). Carbon dioxide production for unamended periphyton (222 mg CO₂-C kg⁻¹ d⁻¹) was significantly greater than unamended detritus (84 mg CO₂-C kg⁻¹ d⁻¹). The response of the heterotrophic microbial community to added C-substrates was related to the nutrient status of the wetland, as substrate-induced respiration (SIR) was higher for detritus than periphyton. Amides and polysaccharides stimulated SIR more than other C-substrates, and methanogenesis was greater contributor to SIR for periphyton than detritus. Inorganic P addition stimulated CO₂ and CH₄ production for periphyton but not detritus, indicating a P limitation in the interior areas of WCA-2a. Continued nutrient loading into oligotrophic areas of WCA-2a or enhanced internal nutrient cycling may stimulate organic matter decomposition and further contribute to undesirable changes to the Everglades ecosystem caused by nutrient enrichment.     

Spatial variation in periphyton fatty acid composition in subtropical streams

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Effects of wood removal on stream habitat and nitrate uptake in two northeastern US headwater streams

Forested headwater streams play an important role in watershed nutrient dynamics, and wood is thought to be a key factor influencing habitat structure and nitrate-nitrogen dynamics in many forested streams. Because wood in streams can promote nitrogen uptake through denitrification, we hypothesized that nitrate uptake velocities would decrease following wood removal. We measured stream characteristics and nitrate uptake velocities before and after wood manipulation experiments conducted at Hubbard Brook Experimental Forest, NH, and the Sleepers River watershed, VT. The mean size of stream substrates and the amount of riffle habitat increased following wood removal. In contrast to our expectations, summer nitrate uptake velocities increased in the wood removal treatments relative to the reference treatments, possibly because wood removal increased the availability of stable substrates for periphyton growth, therefore increasing nitrate demand in these streams. Our results highlight that effects of wood on stream ecosystems occur through multiple pathways and suggest that the relative importance of these pathways may vary seasonally.     

Empirical analysis of the removal rate of periphyton by grazers

To establish a general model for the removal rate of periphyton by grazers, we identified 27 publications in which removal rates could be estimated from grazer enclosure or exclosure experiments. When all the measurements obtained under different experimental conditions were extracted, these publications provided 107 data points. Multiple regression of these data showed that periphyton removal rate increased significantly with grazer body mass and food availability, and decreased with grazer crowding. Grazer body mass explained 65% of the variation, while crowding and food availability explained 7 and 6% respectively. Except for the significantly lower removal rate of amphibians, neither taxon of the grazer nor algal composition significantly affected removal rate. Experiments in the laboratory and in outdoor channels tended to give higher removal rates than experiments performed in streams or lakes. A comparison with previous allometric equations predicting the ingestion rate of other invertebrate guilds, and with experiments in which periphyton ingestion rate was measured as incorporation of labelled food, indicated that a large portion of periphyton is removed by the activity of the grazer rather than by direct ingestion. These results could be utilized to predict the impact of grazing on periphyton biomass.A contribution of the Groupe d'Écologie des eaux douces, Université de Montréal     

Effects of nutrient (N,P, C) enrichment,grazing and depth upon littoral periphyton of a softwater lake

Three field experiments were performed in Lake Lacawac, PA to determine the importance of potentially limiting nutrients relative to other factors (grazing, depth) in structuring shallow water algal periphyton communities. All three experiments measured periphyton growth (as chlorophyll-a, AFDM or biovolumes of the algal taxa) on artificial clay flower pot substrates which released specified nutrients to their outer surfaces.Control of standing crop by nutrient supply rate vs. grazing was examined in Expt. I. Substrates releasing excess N and P, together with one of 4 levels of C (as bicarbonate) were placed either inside or outside exclosures designed to reduce grazer densities. Chlorophyll-a rose from 1.1–25.6 µg.cm^–2, and some dominant taxa (e.g., Oedogonium, Nostoc, Anacystis) were replaced by others (e.g., Scenedesmus, Cryptomonas) as bicarbonate supply increased. Reductions in invertebrate density did not significantly affect chlorophyll-a at any of the nutrient levels.Reasons for the species shift were further evaluated in Expt. II, using a minielectrode to measure the elevation of pH within the periphyton mat through photosynthetic utilization of bicarbonate. The pH adjacent to pots diffusing N, P and large quantities of bicarbonate, and supporting high chlorophyll-a densities of 32 µg cm^–2, averaged 10.0 compared to 6.3 in the water column. Pots diffusing only N and P supported 0.7 µg chlorophyll-a cm^–2 and elevated pH to 8.2. We suspect that bicarbonate addition favored efficient bicarbonate users (e.g., Scenedesmus), while inhibiting other taxa (e.g., Oedogonium) because of the attendant high pH.Expt. III was designed to test effects of depth (0.1 m vs. 0.5 m) and N (NH₄ ⁺ vs. NO₃ ^–) upon the growth response to bicarbonate observed in Expts. I and II. Similar standing crop and species composition were noted on pots at 0.1 m vs. 0.5 m. Enrichment with NH₄ ⁺ vs. NO₃ ^– also appeared to have little effect upon the periphyton community.Shallow water periphyton communities in Lake Lacawac, when supplied with sufficient N and P, appear to show a distinctive response to increasing bicarbonate concentration and pH which is robust to moderate variation in grazer densities, distance from the water surface, and the form of N enrichment.     

Effects of food‐web structure on periphyton stoichiometry in eutrophic lakes: a mesocosm study

1. Aquatic herbivores typically have much higher concentrations of nutrients (e.g. N and P) in their tissues than there is in the food they eat. These stoichiometric differences can cause herbivores to be limited by the elemental quality of their food, which could affect, in turn, the structure of consumer communities and even alter key ecosystem processes. 2. In streams and in the littoral zone of shallow lakes, periphyton is an important food resource for benthic animals. Studying the elemental composition of periphyton may help us to understand food‐web structure, and any reciprocal effect of this structure on periphyton stoichiometry. 3. To understand how alterations in the food‐web structure affect the elemental composition of periphyton in a eutrophic lake, we carried out a long‐term experiment (14 months) in large‐scale mesocosms (40 m³), in which we manipulated food‐web structure, and which were dominated either by planktivorous fish (Rutilus rutilus) or herbivorous invertebrates (without fish). Periphyton was sampled monthly at three depths (0.5, 1.5 and 2.5 m) to determine its biomass and elemental composition (C/N/P ratio). Food‐web structure, physical and chemical parameters were monitored throughout the experiment. 4. Fish had indirect positive effect on periphyton biomass, leading to twofold higher levels than in herbivore‐dominated mesocosms. This result was probably due to control of benthic consumers by fish, suggesting a strong top–down control on periphyton by their consumers in fishless enclosures. 5. The elemental ratios C/P and C/N were lower in deep water in both treatments, mainly mediated by light availability, in accordance with the light/nutrient ratio hypothesis. These ratios were also lower in fishless treatments, probably due to increases in inorganic nutrient availability and grazing pressure in herbivore‐dominated systems. During winter, periphyton elemental composition was similar in both treatments, and was unrelated to inorganic nutrient availability. 6. These results indicate that any alteration of food‐web structure in lakes, such as in biomanipulation experiments, is likely to modify both the biomass and elemental quality of periphyton. Resultant effects on the consumers of periphyton and macrophytes could play a key role in the success of biomanipulations and should be taken into account in further studies.     

ENVIRONMENTAL CONTROL OF STALK LENGTH IN THE BLOOM‐FORMING,FRESHWATER BENTHIC DIATOM DIDYMOSPHENIA GEMINATA (BACILLARIOPHYCEAE)1

Blooms of the freshwater stalked diatom Didymosphenia geminata (Lyngb.) M. Schmidt in A. Schmidt typically occur in oligotrophic, unshaded streams and rivers. Observations that proliferations comprise primarily stalk material composed of extracellular polymeric substances (EPS) led us to ask whether or not the production of excessive EPS is favored under nutrient‐limited, high‐light conditions. We conducted experiments in outdoor flumes colonized by D. geminata using water from the oligotrophic, D. geminata–affected Waitaki River, South Island, New Zealand, to determine the relationship between D. geminata stalk length, cell division rates, and light intensity under ambient and nutrient‐enriched conditions. Stalk lengths were measured in situ, and cell division rates were estimated as the frequency of dividing cells (FDC). FDC responded positively to increasing light intensity and to nutrient additions (N+P and P). Under ambient conditions, stalk length increased as light level increased except at low ambient light levels and temperature. Nutrient enrichment resulted in decreased stalk length and negative correlations with FDC, with this effect most evident under high light. Our results are consistent with the hypothesis that extensive stalk production in D. geminata occurs when cell division rates are nutrient limited and light levels are high. Thus, photosynthetically driven EPS production in the form of stalks, under nutrient‐limited conditions, may explain the development of very high biomass in this species in oligotrophic rivers. The responses of FDC and stalk length under nutrient‐replete conditions are also consistent with occurrences of D. geminata as a nondominant component of mixed periphyton communities in high‐nutrient streams.     

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.