Benthic–pelagic coupling: a comparison of the community structure of benthic and planktonic heterotrophic protists in shallow inlets of the southern Baltic

1. The taxonomic composition, abundance and biomass of heterotrophic protists (ciliates, heterotrophic flagellates (HF), rhizopods and actinopods) in the sediment and water column of shallow inlets of the Southern Baltic was studied under a variety of environmental conditions during 1996–1997. A shallow, highly eutrophic station and a deeper, less eutrophic station were compared.
2. Community biomass ranged from 0.12 to 0.34 μg C cm?3 in the water column and from 1.5 to 105 μg C cm?3 in the sediment. Heterotrophic protists dominated zooplankton biomass at both stations (73% and 84% mean contribution), while they were of minor importance within the zoobenthos. Expressed per unit area, benthic biomass contributed a significant part (44% and 49%) to the total heterotrophic protistan community at both stations.
3. Although the methodology for counting ciliates and HF was focussed on a high taxonomic resolution, the results reveal some general trends in the distribution of heterotrophic protists: protozooplankton biomass was dominated by flagellates (80% mean biomass contribution) at the shallow station and by ciliates (73% mean biomass contribution) at the deep station. In the benthos at both stations, ciliates were the dominant protozoans, followed by the hitherto little‐studied rhizopods (25% and 35% mean biomass contribution) and flagellates.
4. The degree of benthic–pelagic coupling differed between taxonomic groups. Benthic and pelagic communities of ciliates showed little taxonomic overlap. In contrast, many heterotrophic flagellate species were found both in the benthos and in the pelagic. These benthic–pelagic species contributed significantly to the biomass of HF in the water column. The planktonic rhizopod community consisted of a subset of those species found in the benthos.
5. The abundance of benthic and pelagic protists was positively correlated at the shallow station, but taxonomic data indicate that the direct exchange between benthic and pelagic communities was only partly responsible.     

Bacteria and protozoa as integral components of the forest ecosystem--their role in creating a naturally varied soil fertility

The paper explores interactions between the two first organism groups to appear on earth, the bacteria and protozoa, and their interplay with the rest of the ecosystem focusing upon northern boreal forests. The microbial loop is suggested as a mechanism for local inputs of new N to the ecosystem. The possibility to couple short-term microbial processes with their long-term effects, — as registered in plants, soil and the atmosphere, via the abiotic variables — is explored. The latter are investigated in relation to the environments they create for the micro-organisms, and how this results in varying soil fertility. A chain of events is presented that relate high Ca concentration in the mineral soil and high water availability to increased nitrogen availability for plants via the micro-organisms. An example is given of the influence of these parameters directly upon protozoa along an extreme fertility gradient, and also indirect evidence from a Finnish field study of 30 sites with four fertility levels. Finally, there is a discussion about ways to convert knowledge gained in detailed studies of microbial interactions into forms useful when evaluating the present status of and effects of ameliorative management on ecosystems strongly affected by humans. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of dilution stress on the functioning of a saline Mediterranean stream

The effects of seasonality and dilution stress on the functioning of Rambla Salada, a hypersaline Mediterranean stream in SE Spain, were evaluated. The stream is subject to diffuse freshwater inputs from the drainage of intensively irrigated agriculture in the catchment and periodic losses of water through an irrigation channel. Metabolic rates and the biomass of primary producers and consumers were estimated over a 2-year period. During the first year several dilution events occurred, while during the second year the salinity recovery reached predisturbance levels. Functional indicators were compared in the disturbance and recovery salinity periods. Primary production and respiration rates in the Rambla Salada ranged between 0.07–21.05 and 0.19–17.39 g O₂ m⁻² day⁻¹, respectively. The mean values for these variables were 7.35 and 5.48 g O₂ m⁻² day⁻¹, respectively. Mean net daily metabolism rate was 1.87 ± 0.52 g O₂ m⁻² day⁻¹ and mean production/respiration ratio was 2.48 ± 1.1, reflecting autotrophic metabolism. The metabolic rates showed the typical seasonal pattern of Mediterranean open canopy streams. Therefore, gross primary production (GPP) and ecosystem respiration (ER) registered maximum values in summer, intermediate values in spring and autumn and minimum values in winter. The metabolic rates and biomass of consumers were greater in the disturbance period than in the recovery period. However, they did not show significant differences between periods due to their important dependence on seasonal cycle. Seasonality accounted for much of the temporal variability in GPP and ER (76% and 83% in the multiregression models, respectively). Light availability seems to be the most important factor for GPP and ER in the Rambla Salada. Autotrophic biomass responded more to variations in discharge and conductivity than to seasonal variations. In fact, it was severely affected by freshwater inputs after which the epipelic biomass decreased significantly and Cladophora glomerata proliferated rapidly. Epipelic algal biomass was the most sensitive parameter to dilution disturbance. Handling editor: Luigi Naselli-Flores     

Effects of forest fire on headwater stream macroinvertebrate communities in eastern Washington,U.S.A.

1. Recent increases in fire frequency in North America have focused interest on potential effects on adjacent ecosystems, including streams. Headwaters could be particularly affected because of their high connectivity to riparian and downstream aquatic ecosystems through aquatic invertebrate drift and emergence. 2. Headwater streams from replicated burned and control catchments were sampled in 2 years following an intense forest fire in northeastern Washington (U.S.A.). We compared differences in benthic, drift and emergent macroinvertebrate density, biomass and community composition between five burned and five unburned catchments (14–135 ha). 3. There were significantly higher macroinvertebrate densities in burned than control sites for all sample types. Macroinvertebrate biomass was greater at burned sites only from emergence samples; in benthic and drift samples there was no significant difference between burn and control sites. 4. For all sample types, diversity was lower in the burned catchments, and the macroinvertebrate community was dominated by chironomid midges. 5. Compared to the effects of fire in less disturbed ecosystems, this study illustrated that forest fire in a managed forest may have greater effects on headwater macroinvertebrate communities, influencing prey flow to adjacent terrestrial and downstream aquatic habitats for at least the first 2 years post‐fire.     

Meiofaunal responses to nutrient additions in a Mediterranean stream

1. The effects of a moderate addition of nutrients (twofold N and threefold P) were examined during a 2‐year period to determine the response to nutrient addition in a meiofaunal community inhabiting sandy patches in a Mediterranean stream. 2. The pattern of meiofaunal assemblages exhibits a high degree of intra‐ and interannual variability. This pattern alternates between periods of hydrological stability and disturbances, such as floods and droughts, which is a characteristic of Mediterranean systems. 3. A before–after–control–impact (BACI) design was used to determine the outcome of the addition by comparing an upstream non‐enriched reach with an enriched downstream reach. Analysis of the study data by means of a nonparametric permutational procedure (permanova ) showed that fertilisation had a significant effect. Density and biomass values increased in the most abundant meiofaunal groups, including microcrustaceans, oligochaetes and chironomids. Microcrustaceans were the dominant group in the permanent meiofauna. 4. We also examined differences in microcrustacean secondary production in both reaches. Ostracods and cyclopoid copepods increased their secondary production in the impacted reach as a result of the nutrient addition. 5. Our study demonstrated that moderate nutrient enrichment can affect the biomass and production of stream meiofauna, but it is still unclear whether this effect was because of autotrophic or heterotrophic pathways.     

Variable effects of sediment addition on stream benthos

Two upper Piedmont streams were studied to determine the effects of road construction, especially sediment inputs. Benthic macroinvertebrate data suggest that the stream community responded to sediment additions in two different ways. Under high flow conditions the benthic fauna occurs mainly on rocky substrates. As sediment is added to a stream the area of available rock habitat decreases, with a corresponding decrease in benthic density. There is, however, little change in community structure. Under low flow conditions, stable-sand areas may support high densities of certain taxa. Density of the benthic macroinvertebrates in these areas may be much greater than the density recorded in control areas, and there are distinct changes in community structure.     

Leaf litter breakdown in a Mediterranean stream characterised by travertine precipitation

1. Breakdown of four leaf species ( Platanus orientalis , Populus nigra , Salix atrocinerea , Rubus ulmifolius ) was studied in a Mediterranean second-order stream characterised by abundant travertine precipitation, a history of fire in its catchment, and a recently revegetated alluvial corridor.
2. Compared to breakdown rates reported in the literature for congeneric species, breakdown of the four species was slow (k = 0.0024–0.0069 day⁻¹ for the tree species, and 0.0103 and 0.0111 day⁻¹ for Rubus ), in spite of high water temperatures, indicating that the travertine layer that quickly covered submerged leaves impeded decomposer activity and physical fragmentation losses.
3. Breakdown rates nevertheless differed between leaf species in a predictable manner, suggesting that the observed mass loss was largely due to biological processes.
4. The observed tendency towards increasing leaf nitrogen and phosphorus concentrations during breakdown suggests that microorganisms were actively involved in leaf breakdown; however, this interpretation must be viewed with caution because of potentially confounding effects by nutrients contained in the travertine layer.
5. Leaf breakdown of the three indigenous species was faster than that of the exotic species P. orientalis . Due to the recalcitrance of its leaves, the frequent use of Platanus in revegetation schemes following the destruction of indigenous vegetation by fire, exacerbates the negative effect of travertine precipitation on leaf breakdown and, by extension, energy flow in Mediterranean karst streams.     

The new higher level classification of eukaryotes with emphasis on the taxonomy of protists

This revision of the classification of unicellular eukaryotes updates that of Levine et al. (1980) for the protozoa and expands it to include other protists. Whereas the previous revision was primarily to incorporate the results of ultrastructural studies, this revision incorporates results from both ultrastructural research since 1980 and molecular phylogenetic studies. We propose a scheme that is based on nameless ranked systematics. The vocabulary of the taxonomy is updated, particularly to clarify the naming of groups that have been repositioned. We recognize six clusters of eukaryotes that may represent the basic groupings similar to traditional "kingdoms." The multicellular lineages emerged from within monophyletic protist lineages: animals and fungi from Opisthokonta, plants from Archaeplastida, and brown algae from Stramenopiles.     

Effects of sediment releases from a reservoir on stream macroinvertebrates

Effects of sediment release from Guernsey Reservoir on macroinvertebrates of the North Platte River, Wyoming, were investigated during summer 1981. Suspended solids concentrations during sediment release increased from <20 mg l^–1 to >300 mg l^–1. Because fine particulates remained in suspension, mean particle size of substrates was unaltered. Densities of chironomids decreased 90% + during sediment release but recovered to initial levels in 3 weeks after the release ended. Densities of mayflies and oligochaetes increased. Changes in benthic populations were highly correlated with increases in suspended solids.     

Organic matter availability structures microbial biomass and activity in a Mediterranean stream

1. We compared microbial biomass (bacteria, fungi, algae) and the activity of extracellular enzymes used in the decomposition of organic matter (OM) among different benthic substrata (leaves, coarse and fine substrata) over one hydrological year in a Mediterranean stream.
2. Microbial heterotrophic biomass (bacteria plus fungi) was generally higher than autotrophic biomass (algae), except during short periods of high light availability in the spring and winter. During these periods, sources of OM shifted towards autochthonous sources derived mainly from algae, which was demonstrated by high algal biomass and peptidase activity in benthic communities.
3. Heterotrophic activity peaked in the autumn. Bacterial and fungal biomass increased with the decomposition of cellulose and hemicellulose compounds from leaf material. Later, lignin decomposition was stimulated in fine (sand, gravel) and coarse (rocks, boulders and cobbles) substrata by the accumulation of fine detritus.
4. The Mediterranean summer drought provoked an earlier leaf fall. The resumption of the water flow caused the weathering of riparian soils and subsequently a large increase in dissolved organic carbon and nitrate, which led to growth of bacteria and fungi.     

Macrophytic, epipelic and epilithic primary production in a semiarid Mediterranean stream

Summary 1. Primary production by Chara vulgaris and by epipelic and epilithic algal assemblages was measured in a semiarid, Mediterranean stream (Chicamo stream, Murcia, Spain) during one annual cycle. 2. The rates of gross primary production (GPP) and community respiration (CR) were determined for each algal assemblage using oxygen change in chambers. The net daily metabolism (NDM) and the GPPd^?1 : CR₂₄ ratio were estimated by patch‐weighting the assemblage‐level metabolism values. 3. Gross primary production and CR showed significant differences between assemblages and dates. The highest rates were measured in summer and spring, while December was the only month when there were no significant differences in either parameters between assemblages. GPP was strongly correlated with respiration, but not with algal biomass. 4. Chara vulgaris showed the highest mean annual metabolic rates (GPP = 2.80 ± 0.83 gC m^?2 h^?1, CR = 0.76 ± 0.29 gC m^?2 h^?1), followed by the epilithic assemblage (GPP = 1.97 ± 0.73 gC m^?2 h^?1, CR = 0.41 ± 0.12 gC m^?2 h^?1) and epipelic algae (GPP = 1.36 ± 0.22 gC m^?2 h^?1, CR = 0.39 ± 0.06 gC m^?2 h^?1). 5. The epipelic assemblage dominated in terms of biomass (82%) and areal cover (88%), compared with the other primary producers. Epipelic algae contributed 84% of gross primary production and 86% of community respiration in the stream. 6. Mean monthly air temperature was the best single predictor of macrophyte respiration and of epipelic GPP and CR. However, ammonium concentration was the best single predictor of C. vulgaris GPP, and suspended solid concentration of epilithon GPP and CR. 7. Around 70% of the variation in both mean GPP and mean CR was explained by the mean monthly air temperature alone. A multiple regression model that included conductivity, PAR and nitrates in addition to mean monthly air temperature, explained 99.99% of the variation in mean CR. 8. Throughout the year, NDM was positive (mean value 7.03 gC m^?2 day^?1), while the GPP : CR₂₄ ratio was higher than 1, confirming the net autotrophy of the system.     

In situ nutrient assays of periphyton growth in a lowland Costa Rican stream

Nutrient limitation of primary production was experimentally assessed using an in situ bioassay technique in the Quebrada Salto, a third-order tropical stream draining the northern foothills of the Cordillera Central in Costa Rica. Bioassays employed artificial substrata enriched with nutrients that slowly diffuse through an agar-sand matrix (Pringle & Bowers, 1984). Multiple comparisons of regression coefficients, describing chlorophyll-a accrual through time for different nutrient treatments, revealed positive micronutrient effect(s). Micronutrient treatment combinations (Fe, B, Mn, Zn, Co, Mo, EDTA), supplemented with and without nitrate and phosphate, exhibited significantly greater chlorophyll-a accrual over all other treatments (P < 0.05), supporting over three times that of the control after 14-d of substratum colonization. Neither of the major nutrients (N or P) produced a significant stimulation, although the N treatment displayed 50% more chlorophyll-a than the control after 14-d. Similarly, Si, EDTA, and Si + N + P treatments did not exhibit chlorophyll-a response curves that were significantly different from the control. During the experiment, mean NH₄-N and (NO₂ + NO₃)-N concentrations in the Salto were 2.0 µM (28.6 µg · l^–1) and 7.2 µM (100.2 µg · l ^–1), respectively. High concentrations of PO₄-P ( = 2.0 µM; 60.9 µg · l^–1) and TP ( = 3.0 µM; 94.0 µg · l^–1) were also found, and consequently low molar N:P ratios = 4.7). Despite the potential for N limitation in the system, both N and P appear to be at growth saturating levels. This may be due to micronutrient limitation and/or light limitation of periphyton growth in densely shaded upstream portions of the stream.     

Importance of transient storage zones for ammonium and phosphate retention in a sandy-bottom Mediterranean stream

1. The ability of hyporheic sediments to exchange water and retain ammonium and phosphate in the Riera Major stream ,North-East Spain, under different discharge conditions was measured by conducting short-term nutrient and chloride additions. 2. The mean exchange coefficients from free-flowing water to the storage zone (k₁) and vice versa (k₂) were 0.82 × 10–4 s^??1 and 7 × 10^??3 s^??1, respectively. The ratio of storage zone cross-sectional area to stream cross-sectional area (A_S/A) averaged 2.8 × 10–2 and was negatively correlated with discharge (r = –0.85, d.f. = 13, P < 0.001). 3. The percentage of hyporheic zone water which came from surface water varied as a function of discharge and hyporheic depth, ranging between 33% and 95% at 25 cm depth, and between 78% and 100% at 10 cm depth. 4. The nutrient retention efficiency in the hyporheic zone at 10 cm depth measured as uptake length (S_wh) was less than 3.3 cm for ammonium and 37 cm for phosphate. Higher nutrient retentions were measured in the sediments at 10 cm depth than at 25 cm, indicating that near-surface sediments were involved more actively in phosphate retention than the deeper hyporheic sediments. The lack of ammonium at any depth of the hyporheic zone showed that ammonium was very rapidly taken up in the surfacial sediments.     

Controlled release experiments to determine the effects of shade and plants on nutrient retention in a lowland stream

Understanding nutrient uptake and retention in streams remains an important challenge for lotic scientists. In this study a series of pulse and continuous releases of dissolved nutrients were made to shaded and unshaded (reference) reaches of a small lowland stream to determine whether suppression of macrophyte growth by riparian shade impaired nutrient retention. The nutrients were dissolved reactive phosphorus (DRP), total ammoniacal nitrogen (NH₄–N) and nitrate nitrogen (NO₃–N). Nutrient reductions ranged from 100% of DRP when stream water was anoxic, to 5–10% for NH₄–N and NO₃–N in the reference reach. Nutrient removals were affected by travel times in each reach. Percentage removals of NH₄–N (46 ± 10) and NO₃–N (52 ± 14) were higher in the shaded reach than in the swifter moving reference reach (15 ± 8 and 16 ± 10, respectively). DRP (%) removals were 75± 7 and 57 ± 12 for the shaded and reference reaches, respectively. The presence of emergent marginal macrophytes (Persicaria hydropiper) increased stream velocity in the reference reach by reducing the effective channel cross-section area. Shading reduced plant biomass, increased the channel cross-section and lowered velocity in the experimental reach, effecting dramatic reductions in nutrient concentrations over short distances. The opposite effect is more typical for larger, swifter streams having dense stands of submerged macrophytes, where lowering channel plant biomass will cause increased velocities and lower relative nutrient losses. Riparian shade does not necessarily impair nutrient uptake from small streams. Where invasive marginal species such as P. hydropiper dominate headwater streams shade may be beneficial to the protection of downstream waters from eutrophication. Where reduction of nutrient fluxes from small streams is a key objective for protection of downstream waters, active management of streams should seek to increase travel times, allowing greater potential for nutrient uptake. This will need to be weighed against the need for effective drainage in pastoral areas where reduced travel times are usually sought.     

Effects of snow cover on the benthic fauna in a glacier-fed stream

1. Alpine streams above the tree line are covered by snow for 6–9 months a year. However, winter dynamics in these streams are poorly known. The annual patterns of macroinvertebrate assemblages were studied in a glacial stream in the Austrian Alps, providing information on conditions under the snow.
2. Snow cover influenced water temperature, the content of benthic organic matter and insect development. Taxa richness and abundance of macroinvertebrates did not show a pronounced seasonal pattern. The duration of the autumn period with stable stream beds was important in determining the abundance and composition of the winter fauna.
3. There were significant differences in species composition between summer and winter. Two potential strategies in larval survival were evident: adaptation to the extreme abiotic conditions in summer (e.g. Diamesa spp.) or avoidance of these conditions and development during winter (e.g. Ephemeroptera and Plecoptera).
4. A comparison of a stream reach with continuous snow cover and a stream reach that remained open throughout winter showed that conditions under snow are suboptimal. At the open stream site, with higher water temperatures and greater food supply (benthic organic matter content), abundance and taxa richness was higher and larval growth was faster. Several taxa were found exclusively at this site.
5. Winter conditions did not provide an entirely homogeneous environment, abiotic conditions changed rapidly, especially at the onset of snowfall and at snowmelt. Continuous monitoring is necessary to recognize spatial and temporal heterogeneity in winter environments and the fauna of alpine streams.     

Recovery of the macroinvertebrate community below a wastewater treatment plant input in a Mediterranean stream

We sampled chlorophyll a, benthic organic matter, and benthic macroinvertebrates in June 2001 in La Tordera stream (Catalonia, NE Spain), receiving a wastewater treatment plant (WWTP) input. Samples were collected in six equidistant transects in three reaches located upstream (UP), few m below (DW1), and 500 m below the WWTP input (DW2). Our first objective was to assess the effects of the point source on the structure and functional organization of the benthic macroinvertebrate community. Our second objective was to determine if the self-purifying capacity of the stream implied differences between the communities of the DW1 and the DW2 reaches. The WWTP input highly increased discharge, nutrient concentrations, and conductivity and decreased dissolved oxygen. At the DW1 and the DW2 reaches, taxa richness, EPT taxa (Ephemeroptera, Plecoptera, and Trichoptera), and Shannon diversity decreased and gatherer relative density increased relative to the UP reach. At the UP reach, CPOM and FPOM standing crops were similar, whereas at the DW1 and the DW2 reaches CPOM was two times higher than FPOM. Detailed analysis showed that major changes in the benthic community occurred abruptly between 80 and 90 m downstream of the point source (middle of the DW1 reach). At this location, chlorophyll a concentration, density of macroinvertebrates, taxa richness, and scraper relative density increased, whereas gatherer relative percentage decreased. The macroinvertebrate community at the DW2 reach was comparable to that at the second middle of the DW1 reach (DW1B). The macroinvertebrate community at the DW1B and the DW2 reaches were quite similar to that at the UP reach, indicating that the recovery capacity of the stream from nutrient enrichment was high.     

Effects of N and P supply on phytoplankton in Bizerte Lagoon (western Mediterranean)

Enriched bottle experiments were conducted in situ during winter (January and February) and summer (July and August) 2001 to examine the effects of nutrient enrichments (+ N, + P and + NP) on phytoplankton in Bizerte Lagoon, Tunisia. Chlorophyll a (Chl a), ranging from 3.05 μg L⁻¹ in winter to 4.52 μg L⁻¹ in summer, was dominated by the small size-faction (<5 μm) during both seasons. However, the contribution of the large size-fraction (5-200 μm) to Chl a increased from winter (26%) to summer (37%). Similarly, the carbon biomass of the 5-200 μm algae increased during the July/August period that was characterised by the high proliferation of several diatom taxa. In winter, N was the limiting element for phytoplankton growth. Its addition alone (+ N) or with P (+ NP) increased both the <5 μm and 5-200 μm Chl a concentrations. There was no change in the phytoplankton size structure, with the small cells dominating the final algal biomass in all treatments after 5 days. In summer, N and P limited the phytoplankton, but small and large algae exhibited diverse responses to different nutrient enrichments: addition of P increased the Chl a only in the 5-200 μm fraction, the + N treatment enhanced both size classes, and the NP fertilisation mostly stimulated the biomass of large cells. Consequently, the N and P addition in summer was followed by a significant change in the phytoplankton size structure, since both size-fractions contributed equally to the final Chl a biomass. Within the 5-200 μm algal community, various taxa had diverse responses to the nutrient supply during both seasons, leading to a change in the final community composition. The autotrophic flagellates appeared to grow well under N-deficient conditions. In contrast, diatom growth and biomass were mostly stimulated by the N enrichment while dinoflagellates exhibited the highest increase in their growth and biomass with P fertilisation. Our results suggest that the increasing anthropogenic supply of nutrients in the lagoon may influence algal dynamics as well as productivity in different ways depending on the nutrient composition.     

Long‐term moderate nutrient inputs enhance autotrophy in a forested Mediterranean stream

1. The effects of long‐term nutrient addition at moderate levels were examined in the food web of a forested Mediterranean stream. Basal concentrations of N and P were increased twofold (to c. 750 μg N) and threefold (to c. 30 μg P) from ambient concentrations in an experimental reach. Variations in the abundance of microbes (bacteria and algae), meiofauna and macrofauna, microbial processing of organic matter (extracellular enzyme activities) and stoichiometry of biofilms and invertebrates were compared to an upstream control reach during 4 years of artificial nutrient enhancement. 2. Effects were faster in the bacterial compartment but more substantial in the algal compartment. Epilithic algal biomass doubled in the enriched section jointly triggered by nutrients and increased light irradiance in winter and early spring. Only a few animal groups reacted to the enrichment, including the meiofaunal Copepoda, linked to their high use of enriched FPOM, and macrofaunal grazers (Ancylus), which followed the large algal biomass increase. 3. The enrichment caused biofilm phosphatase activity to decrease, while activities related to the use of algal‐related materials (peptidase, β‐glucosidase) increased. Enzymatic activities related to the use of allochthonous organic matter showed only minor and episodic increases. 4. Changes in stoichiometric ratios were apparent in the epilithic compartment, but not in the sand sediment or in the FPOM. Increases in P content were delayed for 9 months in epilithic biofilms and for nearly 2 years in the case of N. 5. After 2 years of enrichment, the flatworm Schmidtea polychroa (predator), oligochaetes (detritivore) and tadpoles of Bufo bufo (grazer) showed higher per cent N. 6. Enrichment effects were produced in spite of flow cessations that occurred commonly in summer. The results show that forested streams subjected to sustained (though minor) nutrient enrichment changed aspects of their biological structure and metabolism and that changes were especially favoured by periods when light was not limiting.