Microbial enzyme activity,nutrient uptake and nutrient limitation in forested streams

1. We measured NH₄⁺ and PO₄^?3 uptake length (S_w), uptake velocity (V_f), uptake rate (U), biofilm respiration and enzyme activity and channel geomorphology in streams draining forested catchments in the northwestern (Northern California Coast Range and Cascade Mountains) and southeastern (Appalachian and Ouachita mountains) regions of the United States. Our goal was to use measures of biofilm enzyme activity and nutrient uptake to assess nutrient limitation in forested streams across broad regional scales. 2. Geomorphological attributes, biofilm enzyme activity and NH₄⁺ uptake were significantly different among streams in the four study units. There was no study unit effect on PO₄^?3 uptake. The proportion of the stream channel in pools, % woody debris, % canopy closure, median substrate size (d₅₀), stream width (w), stream velocity (v), discharge (Q), dispersion coefficient (D) and transient storage (A_s/A) were correlated with biofilm enzyme activity and nutrient uptake in some study units. 3. Canonical correlation analyses across study units revealed significant correlations of NH₄‐V_f and PO₄‐V_f with geomorphological attributes (w, d₅₀, D, % woody debris, channel slope and % pools) and biofilm phosphatase activity. 4. The results did not support our expectation that carbon processing rates by biofilm microbial assemblages would be governed by stream nutrient availability or that resulting biofilm enzyme activity would be an indicator of nutrient uptake. However, the relative abundances of peptidases, phosphatase and glycosidases did yield insight into potential N‐, P‐ and C‐limitation of stream biofilm assemblages, and our use of biofilm enzyme activity represents a novel application for understanding nutrient limitations in forested streams. 5. Regressions of V_f and U against ambient NH₄⁺ and PO₄^?3 indicated that none of our study streams was either NH₄⁺ or PO₄^?3 saturated. The Appalachian, Ouachita and Coastal streams showed evidence of NH₄⁺ limitation; the Ouachita and Coastal streams were PO₄^?3 limited. As a correlate of nutrient limitation and saturation in streams, ratios of total aminopeptidase and phosphatase activities and the ratio of NH₄‐U to PO₄‐U indicate these forested streams are predominantly N‐limited, with only the streams draining Ouachita and Coastal catchments demonstrating appreciable levels of P‐limitation. 6. Our results comparing the stoichiometry of microbial enzyme activity with nutrient uptake ratios and with the molar ratios N and P in stream waters suggest that biological limitations are not strictly the result of stream chemistry and that the assessments of nutrient limitations in stream ecosystems should not be based on chemistry alone. 7. Our present study, along with previous work in streams, rivers and wetlands, suggests that microbial enzyme activities, especially the ratios of total peptidases to phosphatase, are useful indicators of nutrient limitations in aquatic ecosystems.     

Biogeochemical responses of two forest streams to a 2-month calcium addition

1. Calcium (Ca) has been lost from forest soils at the Hubbard Brook Experimental Forest (HBEF) because of decreased atmospheric input of Ca and high input of acid anions. Through time, this Ca loss has led to low streamwater Ca concentration and this change may affect stream ecosystem processes.
2. To test both the biogeochemical response of streams to increased calcium concentration and the role of streams in retaining calcium lost from soils, we added c. 120 μeq Ca L^?1 as CaCl₂ to two second‐order streams at HBEF for 2 months. One stream (buffered) also received an equivalent amount of NaHCO₃ to simulate the increase in pH and alkalinity if Ca were added with associated HCO₃^? ion. The other stream (unbuffered) received only CaCl₂. We collected water samples along a transect above and below the addition site at 11 dates: two before, seven during, and two after the addition.
3. Increase in pH in the buffered stream ranged from 5.6 to about 7.0 in the treated section. There was a net uptake of Ca on all sampling dates during the addition and these uptake rates were positively related to pH. Between 10 and 50% of the added Ca was taken up during the release in the 80‐m study reach. In the unbuffered stream, there was net uptake of Ca on only two dates, suggesting lower Ca uptake.
4. Water samples collected after the addition was stopped showed that a small fraction of the added Ca desorbed from sediments; the remainder was apparently in longer‐term storage in the sediments. No Ca desorbed from the stream sediments in the unbuffered stream, showing that sorption/desorption may be controlled by a pH‐induced increase in the number of exchange sites.
5. These streams appeared to be a significant sink for Ca over a 2‐month time scale, and thus, change in streamwater Ca during a year may be due to processing of Ca within the stream channel, as well as to changes in inputs from the catchment.     

Biogeochemical processes along a nutrient gradient in a tropical ombrotrophic peatland

Biogeochemical properties, including nutrient concentrations, carbon gas fluxes, microbial biomass, and hydrolytic enzyme activities, were determined along a strong nutrient gradient in an ombrotrophic peatland in the Republic of Panama. Total phosphorus in surface peat decreased markedly along a 2.7 km transect from the marginal Raphia taedigera swamp to the interior sawgrass swamp, with similar trends in total nitrogen and potassium. There were parallel changes in the forest structure: basal area decreased dramatically from the margins to the interior, while tree diversity was greatest at sites with extremely low concentrations of readily-exchangeable phosphate. Soil microbial biomass concentrations declined in parallel with nutrient concentrations, although microbes consistently contained a large proportion (up to 47%) of the total soil phosphorus. Microbial C:P and N:P ratios and hydrolytic enzyme activities, including those involved in the cycles of carbon, nitrogen, and phosphorus, increased towards the nutrient-poor wetland interior, indicating strong below-ground nutrient limitation. Soil CO₂ fluxes and CH₄ fluxes did not vary systematically along the nutrient gradient, although potential soil respiration determined on drained soils was lower from nutrient-poor sites. Soil respiration responded strongly to drainage and increased temperature. Taken together, our results demonstrate that nutrient status exerts a strong control on above and below-ground processes in tropical peatlands with implications for carbon dynamics and hence long term development of such ecosystems.     

Trophic state, eutrophication and nutrient criteria in streams

Trophic state is the property of energy availability to the food web and defines the foundation of community integrity and ecosystem function. Describing trophic state in streams requires a stoichiometric (nutrient ratio) approach because carbon input rates are linked to nitrogen and phosphorus supply rates. Light determines the source of carbon. Cross system analyses, small experiments and ecosystem level manipulations have recently advanced knowledge about these linkages, but not to the point of building complex predictive models that predict all effects of nutrient pollution. Species diversity could indicate the natural distribution of stream trophic status over evolutionary time scales. Delineation of factors that control trophic state and relationships with biological community properties allows determination of goals for management of stream biotic integrity.     

Biogeochemical processes in the groundwater discharge zone of urban streams

The influence of biogeochemical processes on nitrogen and organic matter transformation and transport was investigated for two urban streams receiving groundwater discharge during the dry summer baseflow period. A multiple lines of evidence approach involving catchment-, and stream reach-scale investigations were undertaken to describe the factors that influence pore water biogeochemical processes. At the catchment-scale gaining stream reaches were identified from water table mapping and groundwater discharge estimated to be between 0.1 and 0.8 m³ m^?2 d^?1 from baseflow analysis. Sediment temperature profiles also suggested that the high groundwater discharge limited stream water infiltration into the sediments. At the stream reach-scale, dissolved organic carbon (DOC) and dissolved organic nitrogen (DON) concentrations were higher in stream water than in groundwater. However, DOC and DON concentrations were greatest in sediment pore water. This suggests that biodegradation of sediment organic matter contributes dissolved organic matter (DOM) to the streams along with that delivered with groundwater flow. Pore water ammonium (NH₄ ⁺) was closely associated with areas of high pore water DOM concentrations and evidence of sulfate (SO₄ ^2?) reduction (low concentration and SO₄:Cl ratio). This indicates that anoxic DOM mineralization was occurring associated with SO₄ ^2? reduction. However the distribution of anoxic mineralization was limited to the center of the streambed, and was not constrained by the distribution of sediment organic matter which was higher along the banks. Lower sediment temperatures measured along the banks compared to the center suggests, at least qualitatively, that groundwater discharge is higher along the banks. Based on this evidence anoxic mineralization is influenced by groundwater residence time, and is only measurable along the center of the stream where groundwater flux rates are lower. This study therefore shows that the distribution of biogeochemical processes in stream sediments, such as anoxic mineralization, is strongly influenced by both the biogeochemical conditions and pore water residence time.     

Effects of nutrient enrichment on boreal streams: invertebrates, fungi and leaf-litter breakdown

1. The effect of nutrient enrichment on structural (invertebrate indices) and functional (leaf‐litter breakdown rates) characteristics of stream integrity was studied in nine boreal streams. 2. The results showed predicted changes in biotic indices and leaf‐litter breakdown along a complex (principal component) nutrient gradient. Biotic indices were better correlated with nutrient effects than leaf‐litter breakdown. 3. Fungal biomass and invertebrate densities in the litter bags were positively correlated with leaf‐litter breakdown, and both were also positively related to the nutrient gradient. 4. Invertebrate community composition influenced breakdown rate. High breakdown rates at one site were associated with the high abundance of the detritivore Asellus aquaticus. 5. This study lends support to the importance of invertebrate and fungi as mediators of leaf‐litter decomposition. However, our study also shows that study design (length of incubation) can confound the interpretation of nutrient‐induced effects on decomposition.     

Deforestation of watersheds of Panama: nutrient retention and export to streams

A series of eight watersheds on the Pacific coast of Panama where conversion of mature lowland wet forest to pastures by artisanal burning provided watershed-scale experimental units with a wide range of forest cover (23, 29, 47, 56, 66, 73, 73, 91, and 92 %). We used these watersheds as a landscape-scale experiment to assess effects of degree of deforestation on within-watershed retention and hydrological export of atmospheric inputs of nutrients. Retention was estimated by comparing rainfall nutrient concentrations (volume-weighted to allow for evapotranspiration) to concentrations in freshwater reaches of receiving streams. Retention of rain-derived nutrients in these Panama watersheds averaged 77, 85, 80, and 62 % for nitrate, ammonium, dissolved organic N, and phosphate, respectively. Retention of rain-derived inorganic nitrogen, however, depended on watershed cover: retention of nitrate and ammonium in pasture-dominated watersheds was 95 and 98 %, while fully forested watersheds retained 65 and 80 % of atmospheric nitrate and ammonium inputs. Watershed forest cover did not affect retention of dissolved organic nitrogen and phosphate. Exports from more forested watersheds yielded DIN/P near 16, while pasture-dominated watersheds exported N/P near 2. The differences in magnitude of exports and ratios suggest that deforestation in these Panamanian forests results in exports that affect growth of plants and algae in the receiving stream and estuarine ecosystems. Watershed retention of dissolved inorganic nitrogen calculated from wet plus dry atmospheric deposition varied from 90 % in pasture- to 65 % in forest-dominated watersheds, respectively. Discharges of DIN to receiving waters from the watersheds therefore rose from 10 % of atmospheric inputs for pasture-dominated watersheds, to about 35 % of atmospheric inputs for fully forested watersheds. These results from watersheds with no agriculture or urbanization, but different conversion of forest to pasture by burning, show significant, deforestation-dependent retention within tropical watersheds, but also ecologically significant, and deforestation-dependent, exports that are biologically significant because of the paucity of nutrients in receiving tropical stream and coastal waters.     

Initial colonization, nutrient supply, and fungal activity on leaves decaying in streams

Aquatic hyphomycetes dominate leaf decomposition in streams, and their biomass is an important component in the diet of leaf-eating invertebrates. After 2 weeks of exposure in a first-order stream, maple leaf disks had low levels of fungal biomass and species diversity. Spore production by aquatic hyphomycetes also was low. Subsets of these disks were left in the stream for another 3 weeks or incubated in defined mineral solutions with one of three levels of nitrate and phosphate. Stream disks lost mass, increased ergosterol levels and spore production, and were colonized by additional fungal species. External N and P significantly stimulated mass loss, ergosterol accumulation, and spore production of laboratory disks. On disks incubated without added N and P, ergosterol levels declined while conidium production continued, suggesting conversion of existing hyphal biomass to propagules. In all other treatments, approximately equal amounts of newly synthesized biomass were invested in hyphae and conidia. Net yield (fungal biomass per leaf mass lost) varied between 1% (in the laboratory, without added N or P) and 31% (decay in stream). In most treatments, the three aquatic hyphomycete species that dominated spore production during the first 2 weeks in the stream also produced the largest numbers of conidia in the following 3 weeks. Principal-component analysis suggested two divergent trends from the initial fungal community established after 2 weeks in the stream. One culminated in the community of the second phase of stream exposure, and the other culminated in the laboratory treatment with the highest levels of N and P. The results suggest that fungal production in streams, and, by extension, production of invertebrates and higher tropic levels, is stimulated by inorganic N and P.     

Occupancy dynamics of escaped farmed Atlantic salmon in Canadian Pacific coastal salmon streams: implications for sustained invasions

Farmed non-native Atlantic salmon (Salmo salar) is the largest agriculture export product of British Columbia, Canada. Chronic low-volume escapes of salmon from farms into Pacific waters (“leakage”) are typically undetectable (Britton et al. 2011). Analysis of escape-reporting from farmers indicates that reports greatly underestimate the true number of Atlantic salmon inadvertently released from open-net pen rearing sites (Morton and Volpe 2002). To quantify the spatial extent of escaped Atlantic salmon in Canadian Pacific rivers, we systematically snorkel-surveyed 41 known Pacific salmon (Oncorhynchus spp.)-supporting rivers and creeks on Vancouver Island over a span of 3 years. We estimated and accounted for imperfect detections using multi-season occupancy models. We detected Atlantic salmon in 36.6 % of surveyed rivers. After accounting for imperfect detection, occupancy models estimated that over half of surveyed streams across the study area contained Atlantic salmon, and that 97 % of streams with high native salmon diversity were occupied by Atlantic salmon. Even in intensive snorkel surveys, Atlantic salmon are detected in occupied streams only 2/3 the time, suggesting abundance and distribution of non-native salmon is greater than indicated by the only existing data. Further, Atlantic salmon are more likely to occupy streams with high native Pacific salmon diversity—and more likely to maintain occupancy across years—potentially increasing competitive pressure on native salmonids. Understanding local biotic and abiotic predictors of Atlantic salmon occupancy, stream colonization, and local extinction requires more data; the same is true for the effects of escaped Atlantic salmon on local salmon diversity and sustainability. These data for the first time show that Atlantic salmon occupy Pacific coastal rivers for multiple years. The impact of Atlantic salmon occupancy in British Columbia rivers must be factored into policy decisions regarding the future of salmon farming in the provincial waters.     

The effect of salmon redd excavation on stream substrate and benthic community of two salmon spawning streams in Canterbury,New Zealand

The effects of redd excavation by female quinnat salmon (Oncorhynchus tshawytscha walbaum) on the bentic invertebrate communities of two stable springfed spawning streams in the headwaters of the Rakaia River, South Island, New Zealand, were studied during January to September 1982. The gross effects of salmon spawning on the streams appear to be: the loss of instream macrophytes, algae and mosses, a loss of fines and detritus, and a modification of the pool-riffle character. Redd excavation may be associated with a reduction in benthic standing crop of up to 83.6%, with only a minor effect on the benthic community structure. Benthic recolonisation rates were similar for both spawning streams, with almost complete recovery after 95 days.     

Landscape ecotoxicology of coho salmon spawner mortality in urban streams

In the Pacific Northwest of the United States, adult coho salmon (Oncorhynchus kisutch) returning from the ocean to spawn in urban basins of the Puget Sound region have been prematurely dying at high rates (up to 90% of the total runs) for more than a decade. The current weight of evidence indicates that coho deaths are caused by toxic chemical contaminants in land-based runoff to urban streams during the fall spawning season. Non-point source pollution in urban landscapes typically originates from discrete urban and residential land use activities. In the present study we conducted a series of spatial analyses to identify correlations between land use and land cover (roadways, impervious surfaces, forests, etc.) and the magnitude of coho mortality in six streams with different drainage basin characteristics. We found that spawner mortality was most closely and positively correlated with the relative proportion of local roads, impervious surfaces, and commercial property within a basin. These and other correlated variables were used to identify unmonitored basins in the greater Seattle metropolitan area where recurrent coho spawner die-offs may be likely. This predictive map indicates a substantial geographic area of vulnerability for the Puget Sound coho population segment, a species of concern under the U.S. Endangered Species Act. Our spatial risk representation has numerous applications for urban growth management, coho conservation, and basin restoration (e.g., avoiding the unintentional creation of ecological traps). Moreover, the approach and tools are transferable to areas supporting coho throughout western North America.     

Isolation, characterization, and nitrification potential of a methylotroph and two heterotrophic bacteria from a consortium showing methane-dependent nitrification

Abstract A methanotrophic nitrifying consortium was previously obtained from a humisol which showed CH₄-dependent nitrification. Although the methanotroph could not be obtained in pure culture, three other members of the consortium have been isolated: An obligately methylotrophic Methylobacillus (Is-1) which grows only on CH₃OH and does not nitrify; a Pseudomonas (Is-2) which grows on Is-1 culture filtrate and produces NO₂⁻, NO₃⁻ and N₂O from NH₂OH, and NO₃⁻ from NO₂⁻; and a second Pseudomonas (Is-3) which produces NO₃⁻ from NH₄⁺ or NO₂⁻, and N₂O from NH₂OH. A model is proposed for the trophic relations and nitrogen transformations in the consortium which may apply to some natural systems.     

Factors governing the production of juvenile Atlantic salmon in Scottish streams

Data are presented to show how the number and growth of juvenile salmon in streams in the Scottish Highlands are influenced by various physical (temperature, water chemistry, depth and velocity, type of substratum) and biotic (food resources, competition, recruitment) factors.     

Ammonium uptake and mineralization in prairie streams: chamber incubation and short-term nutrient addition experiments

1. We used two separate approaches to estimate ambient ammonium cycling in the north and south branches of Kings Creek, a prairie stream. Chamber experiments were conducted to determine ammonium uptake and mineralization rates associated with epilithic biofilms and filamentous algae collected from the streams. A series of short-term whole-stream ammonium addition experiments were also conducted to estimate the rate of uptake at ambient stream concentrations, based on the relationship between ammonium concentrations and uptake rates.
2. Chamber experiments were scaled up to whole-stream levels, resulting in ambient gross uptake estimates of 0.08  μ g⁻² s⁻¹ for the north branch and 0.16  μ g⁻² s⁻¹ for the south branch. The substrata-specific estimates of mineralization were higher than uptake in both streams.
3. Substrata-specific measurements indicated that ammonium uptake is higher in riffle habitats than in pools habitats. The results of the short-term ammonium addition experiments support these findings.
4. Short-term ammonium addition experiments show that uptake rates saturate with increasing ammonium concentrations. The observed saturation of uptake rates is consistent with a Michaelis–Menten relationship.
5. Scaled estimates of uptake from the chamber experiments were similar to estimates of ambient ammonium uptake based on the whole-stream experiments, and were comparable with previous estimates of ammonium uptake and mineralization made by using stable isotope tracer methods in Kings Creek.     

Effect of periphyton biomass on hydraulic characteristics and nutrient cycling in streams

The effect of periphyton biomass on hydraulic characteristics and nutrient cycling was studied in laboratory streams with and without snail herbivores. Hydraulic characteristics, such as average water velocity, dispersion coefficients, and relative volume of transient storage zones (zones of stationary water), were quantified by performing short-term injections of a conservative tracer and fitting an advection-dispersion model to the conservative tracer concentration profile downstream from the injection site. Nutrient cycling was quantified by measuring two indices: (1) uptake rate of phosphorus from stream water normalized to gross primary production (GPP), a surrogate measure of total P demand, and (2) turnover rate of phosphorus in the periphyton matrix. These measures indicate the importance of internal cycling (within the periphyton matrix) in meeting the P demands of periphyton. Dense growths of filamentous diatoms and blue-green algae accumulated in the streams with no snails (high-biomass streams), whereas the periphyton communities in streams with snails consisted almost entirely of a thin layer of basal cells of Stigeoclonium sp. (low-biomass streams). Dispersion coefficients were significantly greater and transient storage zones were significantly larger in the high-biomass streams compared to the low-biomass streams. Rates of GPP-normalized P uptake from water and rates of P turnover in periphyton were significantly lower in high biomass than in low biomass periphyton communities, suggesting that a greater fraction of the P demand was met by recycling in the high biomass communities. Increases in streamwater P concentration significantly increased GPP-normalized P uptake in high biomass communities, suggesting diffusion limitation of nutrient transfer from stream water to algal cells in these communities. Our results demonstrate that accumulations of periphyton biomass can alter the hydraulic characteristics of streams, particularly by increasing transient storage zones, and can increase internal nutrient cycling. They suggest a close coupling of hydraulic characteristics and nutrient cycling processes in stream ecosystems.     

Modification of benthic insect communities in polluted streams: combined effects of sedimentation and nutrient enrichment

Responses of the benthic insect community of a southern Appalachian trout stream to inorganic sedimentation and nutrient enrichment were monitored over a period of eight months. Entry of pollutants from point sources established differentially polluted zones, allowing an assessment of impacts due to sedimentation alone and in association with elevated nutrient levels. Input of sediment resulted in a significant increase in bed load and decrease of pH at the substrate-water interface (P < 0.05). The zone receiving nutrient runoff from livestock pasture exhibited elevated levels of nitrate and phosphate, but available data indicated such concentrations to be quite low. Species richness, diversity, and total biomass of filter feeding Trichoptera and Diptera, predaceous Plecoptera, and certain Ephemeroptera were significantly reduced in the polluted zones. Inorganic sedimentation, operating indirectly through disruption of feeding and filling of interstitial spaces, was considered to be the primary factor affecting filter feeding taxa. Decomposition of compounds associated with materials in the bed load may depress pH and eliminate acid sensitive species of Plecoptera and Ephemeroptera. Such processes of acidification may be particularly important to Appalachian streams since the pH of regional surface waters is characteristically acidic prior to sedimentation. Accumulation of particles on body surfaces and respiratory structures, perhaps as a function of wax and mucous secretion or surface electrical properties, appears to be the major direct effect of inorganic sedimentation on stream insects. Growths of the filamentous bacterium Sphaerotilus natans were also frequently associated with silted individuals in the zone receiving nutrient addition. Distribution of the bacterium suggested that silted substrates, perhaps as related to the presence of iron compounds, are required for colonization in dilute nutrient solutions. The primary effect of Sphaerotilus colonies appears to be augmentation of particle accumulation through net formation by bacterial filaments. Data indicate that inorganic sedimentation and nutrient addition operate synergistically, eliminating a significantly greater number of taxa than exposure to one pollutant alone.     

The use of periphyton communities for nutrient removal from polluted streams

The results of experiments on the efficiency of periphyton communities for nutrient removal from polluted streams in a continuous flow-through are given. The artifical stream (5 m × 0.7 m × 0.5 m) was made of wood, with silon (a kind of nylon) screens, as a substratum for periphyton growth. The elimination of nutrients was monitored by ammonium, nitrite, nitrate and orthophosphate analyses. In addition, the elimination of organics and the decrease in trophic state were determined. During two field experiments a marked elimination of nitrogen and phosphorus was demonstrated. The maximum efficiency of ammonium and orthophosphate removal was 80% and 70%, respectively. Organic removal reached 35% (C.O.D._Mn-Kubel) and 54% (B.O.D.₅). Inorganic and organic nutrient elimination caused significant changes in periphyton community structure in the outflow portion of the through, evaluated by the saprobic index and the similarity coefficient. The experiments confirmed that periphyton communities are a useful means of nutrient removal from polluted streams.     

Influence of woody debris on nutrient retention in catastrophically disturbed streams

The role of woody debris in nutrient cycling was investigated in two catastrophically disturbed streams in the Pacific Northwest that had been subjected to large inputs of wood. One study site in each catchment had all woody debris removed (take section), while the debris in the other study site was left intact (leave section). Nitrate, phosphate and chloride (a conservative tracer) were released in each section and nutrient retention was monitored at downstream stations. Phosphate was removed from solution more than nitrate, probably due to the high N : P ratio in the stream water. However, there were no major differences in nutrient retention between the take and leave sections. In contrast, experiments in recirculating chambers showed that woody debris and cobbles exhibited higher nitrate and phosphate uptake per unit surface area than sand/gravel or fine particulate organic matter. The high uptake rates of woody debris and cobbles may be related to their suitability for colonization by heterotrophic microorganisms and algae. Wood may not influence nutrient retention significantly at the reach level because of its low surface area relative to other substrates. However, wood may be very important at small spatial scales because of its high uptake activity.     

Initial feeding time of Atlantic salmon,Salmo salar,alevins compared to river flow and water temperature in Norwegian streams

Synopsis The time of initial feeding of Atlantic salmon,Salmo salar, alevins in ten geographically widespread Norwegian streams was estimated theoretically by combining data on spawning time and models describing the time from fertilization to hatching and from hatching to initial feeding at different temperatures. The point of initial feeding was correlated with water flow and water temperature regimes. Initial feeding was avoided in all rivers during spring peak flow, and before water temperature reached 8°C. Two different strategies were indicated: (1) initial feeding may take place before the culmination of the spring flow or (2) after the peak spring flow. The choice of strategy depends on temperature and flow regimes in each river.