Nutrient limitation of algal periphyton in streams along an acid mine drainage gradient

Metal oxyhydroxide precipitates that form from acid mine drainage (AMD) may indirectly limit periphyton by sorbing nutrients, particularly P. We examined effects of nutrient addition on periphytic algal biomass (chl a), community structure, and carbon and nitrogen content along an AMD gradient. Nutrient diffusing substrata with treatments of +P, +NP and control were placed at seven stream sites. Conductivity and SO₄ concentration ranged over an order of magnitude among sites and were used to define the AMD gradient, as they best indicate mine discharge sources of metals that create oxyhydroxide precipitates. Aqueous total phosphorous (TP) ranged from 2 to 23 μg · L^?1 and significantly decreased with increasing SO₄. Mean chl a concentrations at sites ranged from 0.2 to 8.1 μg · cm^?2. Across all sites, algal biomass was significantly higher on +NP than control treatments (Co), and significantly increased with +NP. The degree of nutrient limitation was determined by the increase in chl a concentration on +NP relative to Co (response ratio), which ranged from 0.6 to 9.7. Response to nutrient addition significantly declined with increasing aqueous TP, and significantly increased with increasing SO₄. Thus, nutrient limitation of algal biomass increased with AMD impact, indicating metal oxyhydroxides associated with AMD likely decreased P availability. Algal species composition was significantly affected by site but not nutrient treatment. Percent carbon content of periphyton on the Co significantly increased with AMD impact and corresponded to an increase in the relative abundance of Chlorophytes. Changes in periphyton biomass and cellular nutrient content associated with nutrient limitation in AMD streams may affect higher trophic levels.     

THE RELATIONSHIP OF LIPIDS WITH LIGHT AND CHLOROPHYLL MEASUREMENTS IN FRESHWATER ALGAE AND PERIPHYTON1

Lipid content and lipid class composition were determined in stream periphyton and the filamentous green algae Cladophora sp. and Spirogyra sp, Sterols and phospholipids were compared to chlorophyll a (chl a) as predictors of biomass for stream periphyton and algae. Chlorophyll a, phospholipids, and sterols were each highly correlated with ash-free dry mass (AFDM) (r² > 0.98). Stream periphyton exposed naturally to high light (HL) and low light (LL) had chl a concentrations (μg chl a-mg^?1AFDM) of 7.9± 0.7 and 12.4 ± 2.9, respectively, while the sterol concentrations of these HL and LL stream periphyton (1.6 ± 0.4) were not significantly different (P > 0.05). Periphyton exposed to an irradiance of 300 μmol photons·m^?2s^?1 in the laboratory for 60 h had 5.6 ± 0.55 μg chl a·mg^?1 AFDM, but the same periphyton exposed to 2% incident light for the same amount of time had 11.0 ± 0.56 μg chl a·mg^?1 AFDM. Sterol concentrations in these periphyton communities remained unchanged (1.5 ± 0.3 μg·mg^?1AFDM), Similar results (i.e. changes in chl a but stability of sterol concentrations in response to irradiance changes) were also found for Cladophora and Spirogyra in laboratory experiments. Sterols can be quantified rapidly from a few milligrams of algae and appear to be a useful predictor of eukaryote biomass, whereas cellular levels of chl a vary substantially with light conditions. Phospholipids (or phospholipid fatty acids) are considered to be a reliable measure of viable microbial biomass. Nevertheless, phospholipid content varied substantially and unpredictably among algae and periphyton under different light regimes. Irradiance also had a significant effect on storage lipids: HL Cladophora and HL periphyton had 2 × and 5 × greater concentrations of triacylglycerols, respectively, compared to their LL forms. HL and LL algae also differed in the concentration of several major fatty acids. These light-induced changes in algal lipids and fatty acids have important implications for grazers.     

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.     

Periphyton biomass and ecological stoichiometry in streams within an urban to rural land-use gradient

This study examined the effects land use on biomass and ecological stoichiometry of periphyton in 36 streams in southeastern New York State (USA). We quantified in-stream and land-use variables along a N–S land-use gradient at varying distances from New York City (NYC). Streams draining different landscapes had fundamentally different physical, chemical, and biological properties. Human population density significantly decreased (r = −0.739; P < 0.00001), while % agricultural land significantly increased (r = 0.347; P = 0.0379) with northing. Turbidity, temperature, conductivity, and dissolved Mg, Ca, SRP, pH, DOC, and Si significantly increased in more urban locations, but NO₃ ⁻ and NH₄ ⁺ did vary not significantly along the gradient. Periphyton biomass (as AFDM and Chl-a) in rural streams averaged one-third to one-fifth that measured in urban locations. Periphyton biomass in urban streams averaged 18.8 ± 6.0 g/m² AFDM and 75.6 ± 28.5 mg/m² Chl-a. Urban Chl-a levels ranging between 100 and 200 mg/m², are comparable to quantities measured in polluted agricultural streams in other regions, but in our study area was not correlated with % agricultural land. Periphyton nutrient content also varied widely; algal C varied >20-fold (0.06–1.7 μmol/mm²) while N and P content varied >6-fold among sites. Algal C, N, and P correlated negatively with distance from NYC, suggesting that periphyton in urban streams may provide greater nutrition for benthic consumers. C:N ratios averaged 7.6 among streams, with 91% very close to 7.5, a value suggested as the optimum for algal growth. In contrast, periphyton C:P ratios ranged from 122 to >700 (mean = 248, twice Redfield). Algal-P concentrations were significantly greater in urban streams, but data suggest algal growth was P-limited in most streams regardless of degree of urbanization. GIS models indicate that land-use effects did not easily fit into strict categories, but varied continuously from rural to urban conditions. We propose that the gradient approach is the most effective method to characterize the influence of land use and urbanization on periphyton and stream function.     

USING EPILITHIC ALGAL COMMUNITIES TO ASSESS TROPHIC STATUS IN IRISH LAKES1

Relationships between taxonomic composition of shallow epilithic algal communities and nine environmental variables in 32 lakes of different trophic states in Ireland were explored using gradient analysis. A canonical correspondence analysis using four representative environmental variables, alkalinity (correlated with pH and conductivity), maximum phytoplankton chl a (CHL_max) (correlated with total P, total N, and chl), turbidity, and water color explained 21% of the variance in taxa distributions. The first two axes were significant and accounted for 77% of the variance in the periphyton–environmental relationship. The first axis was strongly related to alkalinity and color, which reflected geology and land use in the watersheds. The second axis was most correlated with CHL_max, and separation of lakes corresponded to their Organization for Economic Cooperation and Development (OECD) trophic classification based on water chemistry. Eutrophic lakes were characterized by cyanobacteria taxa and Stigeoclonium sp. Diatoms and desmids were generally more abundant in oligotrophic and mesotrophic lakes. Values for diatom trophic indices were poor indicators of trophic state. Weighted averaging regression and calibration techniques were used to develop transfer functions between 84 taxa and total P, total N, and CHL_max. The total P inference model predicted OECD trophic classification correctly for 84% of the lakes. Values for taxa preferences resulting from such models can provide the foundation for biomonitoring schemes using extant periphyton communities. The turnover time of periphyton taxa should integrate changes in environmental conditions at a temporal scale intermediate to surface‐sediment fossil diatom assemblages and water column variables, which may be more appropriate for detecting annual changes.     

Periphyton responds differentially to nutrients recycled in dissolved or faecal pellet form by the snail grazer Theodoxus fluviatilis

1. We aimed to separate the effects of grazers on periphyton via grazing from that of nutrient recycling from their faecal pellets. 2. We set up three different experimental treatments (snails/no snails/faecal pellets) and sampled over 16 days. The ‘snail’ treatment contained a low density (snail biomass c. 14 g^?2) of the gastropod grazer Theodoxus fluviatilis and the ‘faecal pellet’ treatment received the same amount of faecal pellets as were produced in the ‘snail’ treatment. Whereas the ‘faecal pellet’ treatment provided extra nutrients to periphyton from the faeces, the ‘snail’ treatment provided nutrients in the form of both faeces and in excreta. There was also direct grazing on periphyton in the ‘snail’ treatment. The ‘no snail’ was not grazed and received no nutrients in faeces or excreta. 3. We measured periphyton C, N and P content, chlorophyll‐a (chl‐a), primary production, bacterial biomass, bacterial production and bacterial respiratory activity. In the water column we measured dissolved inorganic N and soluble reactive P. 4. Snails increased the amount of dissolved inorganic N in the water. On day 16, the periphyton N : P ratio in the ‘faecal pellet’ treatment was lower, and periphyton P content was higher, than in the other two treatments. N : P ratios decreased over time in the ‘faecal pellet’ treatment. Primary and bacterial production were positively correlated in all treatments. 5. Algal chl‐a and primary production of periphyton per unit area and periphyton chl‐a : C ratios increased over the 16 day in the ‘snail’ treatment, and thus excretion of dissolved N by snails had a stronger positive effect on the periphyton community than N and P in faecal pellets. 6. Our data show that excretion and egestion can have different effects on periphyton, probably because of the higher proportion of dissolved N in excreta and the higher proportion of P recycled in faecal pellets. The relative effect of nutrients recycled in egesta or in excretions, probably depends on the form of nutrient limitation of the periphyton. Further, the different components of the periphyton matrix could react differently to the different forms of nutrient recycling. 7. We conclude that direct grazing effects are less important than nutrient effects when nutrients are limiting and grazing pressure is low. Further, the spatial separation of different grazing effects can lead to differences in periphyton production and nutrient stoichiometry. This might be an explanation for the patchiness of periphyton in nature.     

Relationships between benthic biota and hydrological indices in New Zealand streams

1. The objective of this study was to identify the most ecologically relevant hydrological indices for characterizing hydrological regimes in New Zealand streams. To do this we related measures of periphyton chlorophyll a, ash-free dry mass (AFDM), species richness, and diversity and invertebrate density, species richness and diversity, to thirty-four hydrological variables derived from daily flow records at eighty-three sites. The hydrological variables included some describing average flow conditions, flow variability, floods, and low-flow characteristics. 2. A principal components analysis showed that the interrelationship between many of the hydrological variables was high, and most variables correlated significantly with Principal Component 1 (PC1). The flood frequency variables formed a distinct component of the flow regime and were the main contributor to PC2. 3. We found that both the average flow conditions and some measure of variability were significantly related to most of the biological variables, and these individual hydrological variables were more strongly correlated to the biological measures than the composite principal components. Only four of the thirty-four flow variables were significantly correlated (P < 0.05) with measures of periphyton biomass (chlorophyll a and AFDM), whereas twenty-four variables were correlated with periphyton diversity. Conversely, thirty-one of the thirty-four flow variables were correlated with total invertebrate density, whereas only four variables correlated with diversity. 4. We selected the flood frequency (FRE₃), where a flood is defined as flows higher than three times the median flow, as the most ecological useful overall flow variable in New Zealand streams because it explained a significant amount of the variance in four out of the six main benthic community measures, and it had a clear mechanism of control of the biota which was commensurate with current stream ecosystem theory. Periphyton biomass decreased with increasing FRE₃, whereas invertebrate density had an increasing/curvilinear relationship with FRE₃. Periphyton species richness and diversity decreased with increasing FRE₃.     

The effects of hydropsychid colonization on algal response to nutrient enrichment in a small Michigan stream, U.S.A.

1. We tested the hypothesis that the indirect effects of colonization by Hydropsyche spp. (Trichoptera: Hydropsychidae) may be greater than direct effects of nutrients on the benthic algal community growth. Two sets of nutrient-releasing substrates (a total of twenty-four) were deployed into a small pristine stream in northern Michigan. Each set was composed of four treatments replicated three times: (i) no nutrient enrichment (C), (ii) 0.5 M phosphate-P enrichment (P), (iii) 0.5 M nitrate-N enrichment (N) and (iv) 0.5 M phosphate-P plus 0.5 M nitrate-N enrichment (P + N). All hydropsychids colonizing on the substrate in one set (twelve substrates) were removed regularly and the other set (twelve substrates) with undisturbed hydropsychids served as the controls. 2. Algal biomass and gross primary productivity were estimated as chlorophyll a (chl a) concentration, algal biovolume, and carbon fixation rate, respectively. There was a significant interactive effect of hydropsychid colonization and P enrichment on algal biomass measured as chl a concentration. With removal of hydropsychids, chl a concentration increased 11-fold in the P enrichment treatments relative to the controls. The effects of P on chl a was, however, not significant in the presence of hydropsychids. Such interactive effects were not observed when algal responses were measured as biovolume and carbon fixation rate (GPP). 3. It is recommended that algal responses to nutrient enrichment should be measured as biovolume or carbon fixation rate in small streams where hydropsychids are commonly present.     

Effects of upstream lakes and nutrient limitation on periphytic biomass and nitrogen fixation in oligotrophic, subalpine streams

1. We conducted bioassays of nutrient limitation to understand how macronutrients and the position of streams relative to lakes control nitrogen (N₂) fixation and periphytic biomass in three oligotrophic Rocky Mountain catchments. We measured periphytic chlorophyll‐a (chl‐a) and nitrogen‐fixation responses to nitrogen (N) and phosphorus (P) additions using nutrient‐diffusing substrata at 19 stream study sites, located above and below lakes within the study catchments. 2. We found that periphytic chl‐a was significantly co‐limited by N and P at 13 of the 19 sites, with sole limitation by P observed at another four sites, and no nutrient response at the final two sites. On average, the addition of N, P and N + P stimulated chl‐a 35%, 114% and 700% above control values respectively. The addition of P alone stimulated nitrogen fixation by 2500% at five of the 19 sites. The addition of N, either with or without simultaneous P addition, suppressed nitrogen fixation by 73% at nine of the 19 sites. 3. Lake outlet streams were warmer and had higher dissolved organic carbon concentrations than inlet streams and those further upstream, but position relative to lakes did not affect chl‐a and nitrogen fixation in the absence of nutrient additions. Chl‐a response to nutrient additions did not change along the length of the study streams, but nitrogen fixation was suppressed more strongly by N, and stimulated more strongly by P, at lower altitude sites. The responses of chl‐a and nitrogen fixation to nutrients were not affected by location relative to lakes. Some variation in responses to nutrients could be explained by nitrate and/or total N concentration. 4. Periphytic chl‐a and nitrogen fixation were affected by nutrient supply, but responses to nutrients were independent of stream position in the landscape relative to lakes. Understanding interactions between nutrient supply, nitrogen fixation and chl‐a may help predict periphytic responses to future perturbations of oligotrophic streams, such as the deposition of atmospheric N.     

The relationship of floods, drying, flow and light to primary production and producer biomass in a prairie stream

Factors related to autochthonous production were investigated at several sites along a prairie stream at Konza Prairie Research Natural Area. Primary production, algal biomass, litter input, and ability of floods to move native substrate were measured. Additional experiments were conducted to establish the influence of light and water velocity on primary production rates and recovery of biomass following dry periods. The study period encompassed two extreme (> 50 year calculated return time) floods, thus we were able to analyze the effects of scour on periphyton biomass and productivity. Biomass of sedimentary algae was reduced greatly by flooding and did not reach preflood amounts during the 2 months following the first flood. Rates of primary production associated with sediments recovered to levels above preflood rates within 2 weeks. Biomass of epilithic periphyton was not affected as severely as that of sedimentary algae. Little relationship was observed between water velocity and photosythetic rates. Production reached maximum rates at 25% of full sun light. Epilithic chlorophyll levels recovered within eight days following a dry period, and chl a was an order of magnitude greater on rocks than sediments 51 days after re-wetting. Estimated annual rates of primary production were 2.6 times greater in the prairie than in the forest reaches of the stream. The ratio of annual autochthonous:allochthonous carbon input was 4.81 for prairie and 0.32 for the forest. Periphyton production in prairie streams is resilient with regard to flooding and drought and represents a primary carbon source for the system.     

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.     

Sedimentation of chlorophylls in an Arctic fjord under freshwater discharge

Sedimentation of chlorophylls was studied during summer 1997 in Adventfjorden (Spitsbergen, Arctic). During the period of study, the water column was found to be well stratified by a freshened surface layer (salinity <31 PSS). A high load of suspended particulate matter from riverine discharge reduced the euphotic zone to an interval of 0.4–1.1m. Total particulate matter sedimentation rates were about twice as high in June as in July. The following chlorophylls were distinguished in the sedimented particles: chl a and its degradation products (allomer chl a, phaeophytin a, phaeophorbide a, chlorophyllide a), chl b and chl c ₁+c ₂. The quantitatively most important derivative of chl a was phaeophorbide a (31--41% of porphyrin a). Generally, the sedimentation rate of chlorophylls increased with depth. Linear relationships between concentrations of chl a and phaeophorbide a (r ²=0.92), as well as between concentrations of chl a and phaeophytin a (r ²=0.90) indicated a strong connection between phytoplankton abundance and zooplankton grazing. The significant correlation between chl a and chlorophyllide a concentrations (r ²=0.82) showed that most of the sinking chl a belonged primarily to diatoms, and low chlorophyllide a:chl a ratio (0.03) indicated that cellular senescence was not an important reason for the sinking of chl a. Moreover, very low chl b:chl a ratios (about 0.05 calculated for samples where chl b was detectable) suggest that contributions of green algae and/or higher plant detritus were negligible in sinking particles. The ratio of chl c ₁+c ₂:chl a was 0.85 indicating that chl c-containing algae were dominating.     

The role of canopy cover on the recovery of periphyton and macroinvertebrate communities after a month-long flood

We studied the recovery of periphyton and macroinvertebrate communities in a second order stream after a month-long spate that began as increased discharge due to snowmelt in April 2000 but continued through May as a result of frequent rainstorms. We sampled macroinvertebrates and periphyton in June 2000 at three different sites—an upstream site in a conifer tree plantation, a second site ca. 800 m downstream in a mixed hardwood forest, and a third site ca. 1.4 km downstream where there was an open canopy. Periphyton abundance was low on 6 June at all sites but increased 10-fold at the open canopy site the following week (13 June), appearing as a thick mat of Ulothrix. By 26 June, periphyton biomass (as both chlorophyll a and ash free dry mass) had decreased by 80% whereas densities of Chironomidae increased 50-fold and Baetidae increased 4-fold at the open canopy site; little change had occurred in periphyton and macroinvertebrate communities at the forested sites from early to late June. Our results suggest that baetids and chironomids rapidly responded to the increased algal resource at the open canopy site due to their life history characteristics and high growth rates. Chironomids decreased the green mat of Ulothrix by using this alga in tube construction and both chironomids and baetids likely consumed this alga. The varied responses of the periphyton and macroinvertebrate communities at the forested versus open sites demonstrate a resistance to impacts by floods at the forested sites, whereas at the open canopy site periphyton and macroinvertebrate communities were more resilient to the flood disturbance. Handling editor: R. Bailey     

Effects of shading on calcareous benthic periphyton in a short-hydroperiod oligotrophic wetland (Everglades, FL, USA)

The effects of shade on benthic calcareous periphyton were tested in a short-hydroperiod oligotrophic subtropical wetland (freshwater Everglades). The experiment was a split-plot design set in three sites with similar environmental characteristics. At each site, eight randomly selected 1-m² areas were isolated individually in a shade house, which did not spectrally change the incident irradiance but reduced it quantitatively by 0, 30, 50, 60, 70, 80, 90 and 98%. Periphyton mat was sampled monthly under each shade house for a 5 month period while the wetland was flooded. Periphyton was analyzed for thickness, DW, AFDW, chlorophyll a (chl a) and incubated in light and dark BOD bottles at five different irradiances to assess its photosynthesis–irradiance (PI) curve and respiration. The PI curves parameters P _max, I _k and eventually the photoinhibition slope (β) were determined following non-linear regression analyses. Taxonomic composition and total algal biovolume were determined at the end of the experiment. The periphyton composition did not change with shade but the PI curves were significantly affected by it. I _k increased linearly with increasing percent irradiance transmittance (%IT = 1−%shade). P _max could be fitted with a PI curve equation as it increased with %IT and leveled off after 10%IT. For each shade level, the PI curve was used to integrate daily photosynthesis for a day of average irradiance. The daily photosynthesis followed a PI curve equation with the same characteristics as P _max vs. %IT. Thus, periphyton exhibited a high irradiance plasticity under 0–80% shade but could not keep up the same photosynthetic level at higher shade, causing a decrease in daily GPP at 98% shade levels. The plasticity was linked to an increase in the chl a content per cell in the 60–80% shade, while this increase was not observed at lower shade likely because it was too demanding energetically. Thus, chl a is not a good metric for periphyton biomass assessment across variously shaded habitats. It is also hypothesized that irradiance plasticity is linked to photosynthetic coupling between differently comprised algal layers arranged vertically within periphyton mats that have different PI curves.     

Variation in nutrient limitation of lotic and lentic algal communities in a Texas (USA) river

Nutrient limitation of periphyton and phytoplankton was assessed in the Upper Guadalupe River, Texas USA. Nutrient-diffusing substrates with added nitrogen (N) and phosphorus (P) were used to identify the limiting nutrient for lotic algae at three river sites in summer, fall, and winter. Pots enriched with P had significantly higher chlorophyll a concentrations for 7 of 9 trials. Added N alone did not significantly increase algal standing crops, although it was found to be secondarily limiting on one (and possibly two) occasions. Flow-through enrichment experiments were conducted in order to quantify the concentration of P needed to significantly increase algal standing crops. Response to enrichment was rapid when ambient P concentration was low (< 0.010 mg L^–1), but more moderate when ambient P levels were higher (0.015–0.025 mg L^–1). Nutrient limitation of phytoplankton in small surface-release reservoirs varied throughout the study, but N was either primarily or secondarily limiting in 6 of 8 trials; shifts in the limiting nutrient were correlated with fluctuations in flow into the reservoirs. Our enrichment studies show that algal response to nutrient addition was unpredictable as phytoplankton tended to be N-limited while periphyton was mainly P-limited. Further, while discharge apparently dictated the nutrient-biomass relationship for phytoplankton in reservoirs, ambient nutrient level is an important determinant of lotic periphyton response to enrichment.     

Seasonal Dynamics of Benthic and Planktonic Algae in a Nutrient‐Rich Lowland River (Spree,Germany)

We studied chlorophyll a (chl. a), biovolume and species composition of benthic algae and phytoplankton in the eutrophic lower River Spree in 1996. The chl. a concentration was estimated as 3.5 (2.7–4.5) µg/cm² for epipsammon, 9.4 (7.4–11.9) µg/cm² for epipelon and 6.7 (5.7–7.8) µg/cm² for the epilithon (median and 95% C. L.). The mean total biomass of benthic algae was significantly higher (6.0 µg chl. a/cm²) than the areal chl. a content of the pelagic zone (1.6 µg chl. a/cm²). Although certain phytoplankton taxa were abundant in the periphyton, benthic taxa generally dominated the assemblages. Seasonal dynamics of benthic algae were probably controlled by light and nitrate supply (sand), discharge fluctuations (sand, mud) and invertebrate grazing (stones). This paper shows the importance of benthic algae even in phytoplankton‐rich lowland rivers with sandy or muddy sediments. (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Periphyton nutrient status in a temperate stream with mixed land-uses: implications for watershed nitrogen storage

We sampled periphyton communities in a highly productive stream to characterize how longitudinal changes in watershed geology and land use affect periphyton nutrient status and elemental composition. Nutrient status was evaluated from measures of periphyton nutrient composition (carbon, nitrogen, and phosphorus), stable isotope signatures (δ¹⁵N and δ¹³C), and the response of periphyton to experimental enrichment with nitrogen. Biomass and nutrient content increased dramatically from the headwaters to downstream, while tissue nutrient ratios (C:P and C:N) were more consistent and did not indicate strong N- or P-limitation. Nitrogen enrichment experiments did not exhibit a consistent response upstream or downstream, and periphyton C:N:P stoichiometry showed no significant response to N-enrichment. Absolute densities of periphyton N were 5- to 90-fold greater than the overlying N concentrations in stream water (159- to 353-fold greater for P), and the δ¹⁵N signal indicates downstream enrichment from likely watershed sources (urban and agriculture land-use). These results suggest that periphyton in Spring Creek are not N-limited and store large quantities of both N and P, which in turn can be transported downstream during high flow events. Handling editor: David Hamilton     

Reproductive patterns and secondary production of <Emphasis Type="Italic">Gammaropsis japonicus</Emphasis> (Crustacea,Amphipoda) on the seagrass <Emphasis Type="Italic">Zostera marina</Emphasis> of Korea

It is essential to know the nutrient limitation status of biofilms to understand how they may buffer uptake and export of nutrients from polluted watersheds. We tested the effects of nutrient additions on biofilm biomass (chlorophyll a, ash free dry mass (AFDM), and autotrophic index (AI, AFDM/chl a)) and metabolism via nutrient-diffusing substrate bioassays (control, nitrogen (N), phosphorus (P), and N + P treatments) at 11 sites in the Upper Snake River basin (southeast Idaho, USA) that differed in the magnitude and extent of human-caused impacts. Water temperature, turbidity, and dissolved inorganic N concentrations all changed seasonally at the study sites, while turbidity and dissolved inorganic N and P also varied with impact level. Chl a and AI on control treatments suggested that the most heavily impacted sites supported more autotrophic biofilms than less-impacted sites, and that across all sites biofilms were more heterotrophic in autumn than in summer. Nutrient stimulation or suppression of biofilm biomass was observed for chl a in 59% of the experiments and for AFDM in 33%, and the most frequent response noted across all study sites was N limitation. P suppression of chl a was observed only at the most-impacted sites, while AFDM was never suppressed by nutrients. When nutrient additions did have significant effects on metabolism, they were driven by differences in biomass rather than by changes in metabolic rates. Our study demonstrated that biofilms in southeast Idaho rivers were primarily limited by N, but nutrient limitation was more frequent at sites with good water quality than at those with poor water quality. Additionally, heterotrophic and autotrophic biofilm components may respond differently to nutrient enrichment, and nutrient limitation of biofilm biomass should not be considered a surrogate for metabolism in these rivers. Handling editor: D. Ryder     

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.     

Nuisance biomass levels of periphytic algae in streams

Relative coverage of filamentous periphytic algae increased with chlorophyll a (chl a) biomass on natural substrata in 22 northwestern United States and Swedish streams. A biomass range of 100–150 mg chl a m⁻² may represent a critical level for an aesthetic nuisance; below those levels, filamentous coverage was less than 20%. Other indices of water quality (dissolved oxygen content and measures of benthic macroinvertebrate diversity) were apparently unaffected by periphytic biomass or filamentous coverage in these streams. Neither was biomass related to limiting nutrient content (soluble reactive phosphorus, SRP), as has been observed in previous experiments using bare rocks in streams and slides in artificial channels. Ambient SRP concentration may not be a useful predicter of periphyton accrual on natural substrates, due to uptake and recycling of P throughout the stream and undetermined losses such as sloughing and grazing.