The effects of nutrient limitation and stream discharge on the epilithic microbial community in an oligotrophic Arctic stream

Nutrient limitation of epilithic microbial activity is modified by stream discharge and drainage from the tundra surrounding the Kuparuk River, Alaska, USA. During 1984, after three weeks of whole stream enrichment with phosphorus, autotrophic activity per unit biomass had increased in the enriched section of the stream suggesting that phosphorus availability was limiting productivity. In contrast, after three weeks of phosphorus enrichment during 1985, heterotrophic and autotrophic activity was similar in the control and enriched sections of the stream. However, when ammonia or nitrate and phosphorus were added to an in situ bioassay chamber for two weeks, higher community biomass and heterotrophic activity resulted. Ten days later biomass significantly dropped in the unenriched section. Nitrate levels over this period increased four fold concomitantly with decreased stream discharge. Apparently during 1985, nitrogen was limiting epilithic microbial community in the phosphorus enriched section of the Kuparuk River. The significant negative relationship between nitrate concentration and stream discharge observed during 1984 supported the trends seen in 1985. These data suggest that nutrient concentrations which limit epilithic microbial activity and biomass are regulated by the stream discharge and drainage from the surrounding tundra.     

Influence of dissolved organic matter and inorganic nutrients on the biofilm bacterial community on artificial substrates in a northeastern Ohio, USA, stream

Stream bacteria may be influenced by the composition and availability of dissolved organic matter (DOM) and inorganic nutrients, but knowledge about how individual phylogenetic groups in biofilm are affected is still limited. In this study, the influence of DOM and inorganic nutrients on stream biofilm bacteria was examined. Biofilms were developed on artificial substrates (unglazed ceramic tiles) for 21 days in a northeastern Ohio (USA) stream for five consecutive seasons. Then, the developed biofilm assemblages were exposed, in the laboratory, to DOM (glucose, leaf leachate, and algal exudates) and inorganic nutrients (nitrate, phosphate, and nitrate and phosphate in combination) amendments for 6 days. Bacterial numbers in the biofilms were generally higher in response to the DOM treatments than to the inorganic nutrient treatments. There were also apparent seasonal variations in the response patterns of the individual bacterial taxa to the nutrient treatments; an indication that limiting resources to bacteria in stream biofilms may change over time. Overall, in contrast to the other treatments, bacterial abundance was generally highest in response to the low-molecular-weight DOM (i.e., glucose) treatment. These results further suggest that there are interactions among the different bacterial groups in biofilms that are impacted by the associated nutrient dynamics among seasons in stream ecosystems.     

Influence of anthropogenic activities on water quality of a tropical stream ecosystem

Water samples were collected from three sites located in the middle reach of the Njoro River, Kenya, and analysed for total phosphorus (TP), orthophosphate, ammonia‐nitrogen, and nitrate‐nitrogen to evaluate stressor sources (e.g. factories and wastewater ponds) and the general stream water quality. The stream surface water was also analysed for biochemical oxygen demand (BOD₅) to provide an overview of organic matter loading. Mugo, Egerton Bridge and the canning factory sites of the Njoro River had low water quality which is likely to be due to poor farming, partially treated effluents and poor provision of sanitation facilities to the riparian communities. The concentrations of the selected nutrients did not differ significantly among the three sites, presumably due to pollution of the whole stream reach by the catchment nutrient sources. High phosphate concentrations (i.e. ～0.76 mgPO₄ l^?1 and ～0.87 mgTP l^?1) at Canning Factory were recorded during the low flow dry season. Nitrate‐nitrogen concentrations varied significantly with water discharge which explained between 63 and 87% of the nutrient variability in the three sites. BOD₅ differed significantly among the three sites, with historical effects of wastewater and factory effluent discharge being reflected in the results of Egerton Bridge and Canning Factory. The concentrations of ammonia‐nitrogen, TP and orthophosphate were higher in the wastewater than in the river water whereas nitrate‐nitrogen was lower. This study indicates that the Njoro River is stressed by nutrients from the activities within its catchment. With the increasing population, the nutrient load to the river will continue to increase and the water quality will continue to deteriorate. Reductions of nutrient loads into the river as well as provision of sanitation facilities to the riparian communities are needed to control further water degradation.     

Characterization and qualitative changes in DOM chemical characteristics related to hydrologic conditions in a Pampean stream

The aim of our study was; (i) to characterize the composition of DOM in stream water and their potential sources (groundwater, overland flow, subsurface flow and rain water) and (ii) to analyze changes in DOM concentration and composition under different hydrological conditions (baseflow and high flow) in a third-order Pampean stream (Argentina). Pampean streams are mainly fed by the shallow aquifer under baseflow conditions and they lack of riparian forest. In addition, water velocity is low due to the gentle slope of the region and nutrient levels are high, favoring the development of rich macrophyte communities. DOM optical properties in the stream and end members were determined by combining absorbance-fluorescence spectroscopy techniques. Our results indicated that DOM chemical characteristics in the stream were mainly modulated by a differential contribution of end members to stream water depending on hydrological conditions. We observed that DOM in groundwater showed a microbial origin while DOM in runoff was terrestrially-derived. DOC concentration and inputs of humic substances from the riparian zone increased with discharge at high flow conditions. Due to the strong link between DOC properties and the riparian environment, structural alterations in the stream channel and changes in riparian vegetation (forestation) may result in changes in DOM composition and dynamics.     

Microbial leaf degraders in boreal streams: bringing together stochastic and deterministic regulators of community composition

1. Leaves that fall into the water represent a new habitat for microorganisms to colonise in streams, providing an opportunity to study colonisation and the subsequent regulation of community structure. We explored community composition of bacteria and fungi on decomposing alder leaves in nine streams in central Sweden, and describe their relationship with environmental variables. Succession of the microbial community was studied in one of the streams for 118 days. Microbial community composition was examined by denaturing gradient gel electrophoresis on replicate samples of leaves from each stream. 2. During succession in one stream, maximum taxon richness was reached after 34 days for bacteria and 20 days for fungi respectively. Replicate samples within this stream differed between each other earlier in colonisation, while subsequently such variation among replicate communities was low and remained stable for several weeks. Replicate samples taken from all the nine streams after 34 days of succession showed striking similarities in microbial communities within‐streams, although communities differed more strongly between streams. 3. Canonical analysis of microbial communities and environmental variables revealed that water chemistry had a significant influence on community composition. This influence was superimposed on a statistical relationship between the properties of stream catchments and microbial community composition. 4. The catchment regulates microbial communities in two different ways. It harbours the species pool from which the in‐stream microbial community is drawn and it governs stream chemistry and the composition of organic substrates that further shape the communities. We suggest that there is a random element to colonisation early in succession, whereas other factors such as species interactions, stream chemistry and organic substrate properties, result in a more deterministic regulation of communities during later stages.     

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

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

Extracellular phosphatase activity in sediments of the Breitenbach,a Central European mountain stream

Activity of extracellular phosphatases (phosphomonoesterases) was measured in sandy streambed sediments of the Breitenbach, a small unpolluted upland stream in Central Germany. Fluorigenic 4-methylumbelliferyl phosphate served as a model substrate. Experiments were conducted using sediment cores in a laboratory simulation of diffuse groundwater discharge through the stream bed, a natural process occurring in the Breitenbach as well as many other streams.Streambed sediments contained high levels of particulate phosphorus, but concentrations of dissolved phosphorus in the interstitial water were 3 to 4 orders of magnitude lower. These interstitial concentrations were similar to those in the stream and groundwater. Extracellular phosphatase activity was high in the streambed sediments. These enzymes probably contribute significantly to the flux of phosphorus in sediment by hydrolyzing phosphomonoesters, making free phosphate available to the sediment microorganisms.Factors influencing the kinetic parameters V _max (maximum activity) and apparent K _m (enzyme affinity) of phosphatase were discharge rates of water through the sediment, water quality (ground- or stream water), and substrate (phosphomonoesters) as well as dissolved ortho-phosphate concentrations. Enzymes are supposed to be effective at limiting substrate concentrations, where, in this study, changes in discharge rates had little influence on rates of hydrolysis. Higher V _max and lower K _m values were found during percolation of groundwater through the sediment cores, compared with stream water. This indicates that rates of hydrolysis were higher with groundwater, both at substrate limitation and at substrate saturation. This was probably a consequence of the lower levels of dissolved ortho-phosphate in the groundwater.     

Contrasting nutrient exports from a forested and an agricultural catchment in south-eastern Australia

Dissolved organic carbon (DOC) and total and inorganic nitrogen and phosphorus concentrations were determined over 3 years in headwater streams draining two adjacent catchments. The catchments are currently under different land use; pasture/grazing vs plantation forestry. The objectives of the work were to quantify C and nutrient export from these landuses and elucidate the factors regulating export. In both catchments, stream water dissolved inorganic nutrient concentrations exhibited strong seasonal variations. Concentrations were highest during runoff events in late summer and autumn and rapidly declined as discharge increased during winter and spring. The annual variation of stream water N and P concentrations indicated that these nutrients accumulated in the catchments during dry summer periods and were flushed to the streams during autumn storm events. By contrast, stream water DOC concentrations did not exhibit seasonal variation. Higher DOC and NO₃ ⁻ concentrations were observed in the stream of the forest catchment, reflecting greater input and subsequent breakdown of leaf-litter in the forest catchment. Annual export of DOC was lower from the forested catchment due to the reduced discharge from this catchment. In contrast however, annual export of nitrate was higher from the forest catchment suggesting that there was an additional NO₃ ⁻ source or reduction of a NO₃ ⁻ sink. We hypothesize that the denitrification capacity of the forested catchment has been significantly reduced as a consequence of increased evapotranspiration and subsequent decrease in streamflow and associated reduction in the near stream saturated area.     

Mathematical modelling as a tool for management in eutrophication control of shallow lakes

The eutrophication model Delwaq-Bloom-Switch is developed to be a functional tool for water management. Therefore it includes nutrients, algal biomass and composition as well as water transparency. A module describing the interaction between water and bottom gives the model the flexibility to deal with measures, such as a decrease of the external phosphorus loading and flushing with water differing in composition from the lake water. This paper focuses on the functional aspects of the model, the results of an application on Lake Veluwe, The Netherlands, and the implications for water management.With one set of coefficients DBS reproduces the most important characteristics of Lake Veluwe for a period of two years before measures (reduction of the external loading and flushing during the winter months) and eight years after the measures. The phosphorus concentration decreased and became growth limiting for algae instead of nitrogen and light. Both in measurements and modelling results, the algal composition changed from blue-green algae dominance to green algae and diatom dominance. Lake Veluwe had a relatively short transient phase after reduction of external loading, because high nitrate concentrations in the flushing water inhibited a long period with high phosphorus releases from the bottom.Model calculations were carried out to investigate the effects of fish stock management and optimization of flushing. Both measures are promising.     

Interactive effects of nutrients and contaminants from pulp mill effluents on riverine benthos

Bleached kraft mill effluents (BKME) contain both nutrients(nitrogen and phosphorus) and contaminants that can have a stimulatoryor inhibitory effect on riverine food webs. For the Northern RiverBasins Study (NRBS), separation of the effects of nutrients andcontaminants in BKME was an important issue because field biomonitoringwas unable to isolate these impacts. We addressed this problem by usingriverside mesocosm experiments, along with field observations, todetermine the effects of BKME on benthic food webs of the upperAthabasca River near Hinton, Alberta, Canada. Response variablesincluded periphyton biomass, benthic insect density and composition, andthe biomass of abundant invertebrates (i.e., caddisfly, mayfly andstonefly taxa). Experiments were conducted during autumn when nutrientenhancement was hypothesized to increase autochthonous primaryproduction as a result of low river discharge and high water clarity.Treatments included reference river water, 1% BKMEconcentrations, and nutrients (nitrogen plus phosphorus) equivalent tothe 1% BKME treatment. Although pulp mill effluents can inducesublethal toxicity in benthic biota, BKME concentrations equivalent tolevels in the Athabasca River did not cause measurable toxicity.Relative to reference water treatments, BKME-associated increases inavailable phosphorus produced enrichment effects including increases inperiphyton and insect biomass, and invertebrate abundance. Insect familyrichness was not affected by the treatments. The development andapplication of a field-based artificial stream system for the NRBSimproved our mechanistic understanding of the effects of BKME on benthicbiota of nutrient and contaminant stressors. By combining artificialstream results with field observations, we were also able to link thismechanistic understanding of stressor effects directly to impacts in theriver. We conclude that the response to BKME in the Peace-AthabascaBasin is largely one of nutrient enrichment rather than that oftoxicity.     

Shape variation in a benthic stream fish across flow regimes

Evolution of fish body shapes in flowing and non-flowing waters have been examined for several species. Flowing water can select for fish body shapes that increase steady swimming efficiency, whereas non-flowing water can favor shapes that increase unsteady swimming efficiency. Benthic stream fishes often use areas near the substrate that exhibit reduced or turbulent flow, thus it is unclear which swimming forms would be favored in such environments, and how shape might change across flow regimes. To test the relationship between fish body shape and flow regime in a benthic stream fish, we used geometric morphometric techniques to characterize lateral body shape in mountain sucker (Catostomus platyrhynchus) across flow rates, using stream gradient as an indicator of stream flow. Mountain suckers from low-flow environments were more streamlined, consistent with steady swimming body shapes, whereas mountain suckers from high flows had deeper bodies, consistent with unsteady swimming body shapes. In addition, smaller individuals tended to have more robust body shapes. These patterns are opposite to those predicted for stream fishes in the mid-water column. The benthic stream environment represents a distinct selective environment for fish shape that does not appear to conform to the simple dichotomy of flowing versus non-flowing water.     

The effects of storms,heavy rain,and sedimentation on the shallow coral reefs of St. John,US Virgin Islands

Efforts to limit plant growth in streams by reducing nutrients would benefit from an understanding of the relative influences of nutrients, streamflow, light, and other potentially important factors. We measured macrophytes, benthic algae, nutrients in water and sediment, discharge, and shading from 30 spring-fed or runoff-influenced streams in the upper Snake River basin, ID, USA. We hypothesized that in hydrologically stable, spring-fed streams with clear water, macrophyte and benthic algae biomass would be a function of bioavailable nutrients in water or sediments, whereas in hydrologically dynamic, runoff-influenced streams, macrophyte and benthic algae biomass would further be constrained by flow disturbance and light. These hypotheses were only partly supported. Nitrogen, both in sediment and water, was positively correlated with macrophyte biomass, as was loosely sorbed phosphorus (P) in sediment. However, P in water was not. Factors other than nutrient enrichment had the strongest influences on macrophyte species composition. Benthic algal biomass was positively correlated with loosely sorbed sediment P, lack of shade, antecedent water temperatures, and bicarbonate. These findings support the measurement of bioavailable P fractions in sediment and flow histories in streams, but caution against relying on macrophyte species composition or P in water in nutrient management strategies for macrophytes in streams.     

Nutrient limitation of phytoplankton in a central Arizona reservoir

The influence of added nutrients nitrogen, phosphorus and carbon on the phytoplankton of a small recreational reservoir in central Arizona was investigated during the summer, 1974. Polyethylene bags were used to isolate lake water and the natural populations for the addition of nitrogen, phosphorus and carbon individually and in combination. A large increase in phytoplankton numbers, extractable chlorophyll, pH and dissolved oxygen occurred only in bags to which both nitrogen and phosphorus were added, suggesting that both nitrogen and phosphorus levels were limiting to the primary producers. Little alteration in species composition resulted from the addition of the above nutrients.     

Nutrient and Sediment Retention in a Temperate Marsh Ecosystem

The Wye marsh, an undeveloped wetland of 639.9 ha, serves as a receiving water for a 19,600 ha agricultural watershed before discharging into the Great Lakes. Seasonal and diel changes in limnological variables, most notably nitrogen, phosphorus, silica and turbidity, have been examined in the Wye river, marsh and discharge waters over an 11-month period and related to the composition of marsh sediments. Retention rates of the marsh ecosystem for incoming nutrients and suspended solids have been estimated by calculations of mean monthly inflow to outflow ratios using the levels of these factors in the Wye River and the marsh discharge. Calculated mean monthly retention rates of the marsh for incoming nutrients is estimated at no less than 61% for nitrogen-N, 36% for total phosphate, and 14% for soluble silica during the ice-free period. Based on observed turbidity levels, the retention of incoming suspended solids is estimated at 65% for the same period. The Wye marsh is a substantial sump for inorganic and organic materials which are retained within the water column, biomass and unconsolidated sediments.     

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

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

Geomorphology variables predict fish assemblages for forested and endorheic rivers of two continents

Stream fishes are restricted to specific environments with appropriate habitats for feeding and reproduction. Interactions between streams and surrounding landscapes influence the availability and type of fish habitat, nutrient concentrations, suspended solids, and substrate composition. Valley width and gradient are geomorphological variables that influence the frequency and intensity that a stream interacts with the surrounding landscape. For example, in constrained valleys, canyon walls are steeply sloped and valleys are narrow, limiting the movement of water into riparian zones. Wide valleys have long, flat floodplains that are inundated with high discharge. We tested for differences in fish assemblages with geomorphology variation among stream sites. We selected rivers in similar forested and endorheic ecoregion types of the United States and Mongolia. Sites where we collected were defined as geomorphologically unique river segments (i.e., functional process zones; FPZs) using an automated ArcGIS‐based tool. This tool extracts geomorphic variables at the valley and catchment scales and uses them to cluster stream segments based on their similarity. We collected a representative fish sample from replicates of FPZs. Then, we used constrained ordinations to determine whether river geomorphology could predict fish assemblage variation. Our constrained ordination approach using geomorphology to predict fish assemblages resulted in significance using fish taxonomy and traits in several watersheds. The watersheds where constrained ordinations were not successful were next analyzed with unconstrained ordinations to examine patterns among fish taxonomy and traits with geomorphology variables. Common geomorphology variables as predictors for taxonomic fish assemblages were river gradient, valley width, and valley slope. Significant geomorphology predictors of functional traits were valley width‐to‐floor width ratio, elevation, gradient, and channel sinuosity. These results provide evidence that fish assemblages respond similarly and strongly to geomorphic variables on two continents.     

Nutrient pulsing as a regulator of phytoplankton abundance and community composition in Galveston Bay, Texas

Galveston Bay, Texas, is a large shallow estuary with a watershed that includes 60% of the major industrial facilities of Texas. However, the system exhibits low to moderate (2-20 μg l⁻¹) microalgal biomass with sporadic phytoplankton blooms. Both nitrogen (N) and phosphate (P) limitation of phytoplankton growth have been proposed for the estuary. However, shifts between N and P limitation of algae growth may occur due to annual fluctuations in nutrient concentrations. The primary goal of this work was to determine the primary limiting nutrient for phytoplankton in Galveston Bay. Nutrient addition bioassays were used to assess short-term (1-2 days) phytoplankton responses (both biomass and community composition) to potentially limiting nutrients. The experimental bioassays were conducted over an annual cycle using natural water collected from the center to lower part of the estuary. Total phytoplankton biomass increased in the nitrate (10 μM) additions in 11 of the 13 bioassays, but no significant increases were detected in the phosphate (3 μM)-only additions. Bioassay results suggest that the phytoplankton community was usually not phosphate limited. All major groups increased in biomass following nitrate additions but diatoms increased in biomass at a faster rate than other groups, shifting the community composition toward higher relative abundance of diatoms. The results of this study suggest that pulsed N input events preferentially favor increases in diatom biomass in this estuary. The broader implications of this study are that N pulsing events, primarily due to river discharge, play an important role in structuring the phytoplankton community in the Galveston Bay estuary.     

Validation of the ion-exchange membrane bioreactor concept in a plate-and-frame module configuration

The ion exchange membrane bioreactor (IEMB) is a particular case of a membrane-supported biofilm reactor, in which oxy-anions, used as electron acceptors by an anoxic mixed microbial culture, are removed from a polluted water stream through an anion-exchange membrane. The opposite side of this membrane is used for the development of a biofilm, contacting a biocompartment, to which nutrients and chloride are fed as a source of “driving” counter-ion. The applicability of a plate-and-frame IEMB module configuration, consisting of a series of membranes, for the treatment of drinking water contaminated with nitrate and perchlorate, was evaluated. Permeation of carbon source across the membrane to the treated water stream was avoided by a dedicated start-up procedure involving a gradual increase of ethanol feeding to the IEMB biocompartment. It was demonstrated that the biocompartment pH must be controlled not only to guarantee a complete perchlorate removal, but also to avoid precipitation of struvite on the membrane surface, which provokes membrane scaling and decreases the availability of nutrients for the biofilm. Under these conditions, the IEMB was successfully operated maintaining both nitrate and perchlorate concentrations in the treated water below their recommended levels for drinking water supplies.     

Limiting nutrients for periphyton growth in sub-alpine, forest, agricultural and urban streams

1. Nutrient-diffusing artificial substrata were used in summer and autumn to assess limiting nutrients for periphytic algal growth in streams draining sub-alpine, forested, agricultural and urban catchments in eastern Victoria, Australia. 2. Chlorophyll a density was primarily limited by nitrogen in most cases; often phosphorus was secondarily limiting. One sub-alpine, one forest and one agricultural stream were primarily phosphorus limited in at least one season. Added trace elements and vitamins did not increase chlorophyll density. 3. The dominant filamentous algal genera did not differ greatly between various nutrient enrichments. However, the relative abundance of Stigeodanium spp. was sometimes increased on substrata containing the limiting nutrient. 4. The results suggest that nutrient limitation is a widespread phenomenon in Victorian streams, and that limiting nutrients can be inferred from stream-water nitrogen to phosphorus ratios in many instances.