Whole-stream nitrate addition affects litter decomposition and associated fungi but not invertebrates

We assessed the effect of whole-stream nitrate enrichment on decomposition of three substrates differing in nutrient quality (alder and oak leaves and balsa veneers) and associated fungi and invertebrates. During the 3-month nitrate enrichment of a headwater stream in central Portugal, litter was incubated in the reference site (mean NO₃-N 82 μg l⁻¹) and four enriched sites along the nitrate gradient (214–983 μg NO₃-N l⁻¹). A similar decomposition experiment was also carried out in the same sites at ambient nutrient conditions the following year (33–104 μg NO₃-N l⁻¹). Decomposition rates and sporulation of aquatic hyphomycetes associated with litter were determined in both experiments, whereas N and P content of litter, associated fungal biomass and invertebrates were followed only during the nitrate addition experiment. Nitrate enrichment stimulated decomposition of oak leaves and balsa veneers, fungal biomass accrual on alder leaves and balsa veneers and sporulation of aquatic hyphomycetes on all substrates. Nitrate concentration in stream water showed a strong asymptotic relationship (Michaelis–Menten-type saturation model) with temperature-adjusted decomposition rates and percentage initial litter mass converted into aquatic hyphomycete conidia for all substrates. Fungal communities did not differ significantly among sites but some species showed substrate preferences. Nevertheless, certain species were sensitive to nitrogen concentration in water by increasing or decreasing their sporulation rate accordingly. N and P content of litter and abundances or richness of litter-associated invertebrates were not affected by nitrate addition. It appears that microbial nitrogen demands can be met at relatively low levels of dissolved nitrate, suggesting that even minor increases in nitrogen in streams due to, e.g., anthropogenic eutrophication may lead to significant shifts in microbial dynamics and ecosystem functioning. Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Effect of nutrients on the sporulation and diversity of aquatic hyphomycetes on submerged substrata in a Mediterranean stream

Eutrophication in streams often results in enhanced decomposition rates and chemical conditions that greatly affect the suitability of the habitat for many species. We performed an enrichment experiment in a 50-m reach of a Mediterranean stream by the continuous addition of N and P (enriched reach) while another consecutive reach upstream served as the reference (unenriched reach). After 1.5 years of enrichment, conidia production and species composition of aquatic hyphomycetes were measured in communities colonizing four streambed substrata (Platanus acerifolia and Populus nigra leaves, epipsammic and epilithic biofilms). Samples collected from the two reaches were subjected to two nutrient treatments in the laboratory [low nutrient (−NP) and high nutrient (+NP) concentrations] to test for short-term (48 h) nutrient effects. Long-term nutrient additions did not affect potential sporulation rates while short-term nutrient enrichment produced insignificant increase in these rates on most substrata collected in the unenriched reach. Major differences in potential sporulation rates were observed between substrata, the highest rate being for leaves (especially Platanus acerifolia). However, enrichment produced significant changes in fungal community composition. Clavariopsis aquatica, Alatospora acuminata and Lemonniera sp. were dominant in the enriched reach while Heliscella stellata, Trisceloporus acuminatus and Clavatospora longibrachiata were characteristic of the unenriched reach. Our results suggest that fungal N demands can be fulfilled at relatively low levels of dissolved nitrate and further increases in the nutrients availability may not result in enhanced fungal activity.     

Effects of nutrient enrichment and leaf quality on the breakdown of leaves in a hardwater stream

1. The breakdown of leaf litter in streams is influenced strongly by leaf quality and the concentration of dissolved nutrients, primarily inorganic nitrogen (N) and phosphorus (P) in the water. We examined the effect of nutrient enrichment on the breakdown of three species of leaves in a hardwater, nutrient‐rich stream. The rate of microbial respiration was also measured on the decomposing leaves. 2. The breakdown rates of dogwood (Cornus stolonifera), aspen (Populus tremuloides) and birch (Betula occidentalis), k‐values of 0.0461, 0.0307 and 0.0186 day^–1, respectively, were unaffected by nutrient enrichment and generally faster than reported previously. Microbial respiration on the leaves was greater than reported previously for leaves of congeneric species. It appears that leaf breakdown in the study stream was not nutrient limited. 3. Nitrogen‐based measures of leaf quality, such as percentage N and carbon (C)/nitrogen ratio, did not correspond to measured breakdown rates among the three leaf types. The best predictors of relative breakdown rates were percentage lignin and the percentage of the total carbon that occurred as lignin. We suggest that, when leaf breakdown is not nutrient limited, measures of carbon quality (i.e. lignin‐based measures) are a better assessment of overall leaf quality than are N‐based measures. 4. Previous studies have indicated that the enzymes produced by aquatic hyphomycetes (microfungi) operate most efficiently at a basic pH and in the presence of calcium ions. The hardwater conditions (pH=8.6, total hardness > 300 mg CaCO₃ L^–1) and abundance of dissolved NO₃ and soluble reactive phosphorous (SRP) (approximately 50 μg L^–1, each) in the study stream appear to have provided conditions that resulted in a high respiration rate and rapid breakdown of leaf litter.     

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.     

Elevated temperature may intensify the positive effects of nutrients on microbial decomposition in streams

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Response of Periphytic Algae to Gradients in Nitrogen and Phosphorus in Streamside Mesocosms

In this study we manipulated both nitrogen and phosphorus concentrations in stream mesocosms to develop quantitative relationships between periphytic algal growth rates and peak biomass with inorganic N and P concentrations. Stream water from Harts Run, a 2nd order stream in a pristine catchment, was constantly added to 36 stream-side stream mesocosms in low volumes and then recirculated to reduce nutrient concentrations. Clay tiles were colonized with periphyton in the mesocosms. Nutrients were added to create P and N concentrations ranging from less than Harts Run concentrations to 128 μg SRP l⁻¹ and 1024 μg NO₃-N l⁻¹. Algae and water were sampled every 3 days during colonization until periphyton communities reached peak biomass and then sloughed. Nutrient depletion was substantial in the mesocosms. Algae accumulated in all streams, even streams in which no nutrients were added. Nutrient limitation of algal growth and peak biomass accrual was observed in both low P and low N conditions. The Monod model best explained relationships between P and N concentrations and algal growth and peak biomass. Algal growth was 90% of maximum rates or higher in nutrient concentrations 16 μg SRP l⁻¹ and 86 μg DIN l⁻¹. These saturating concentrations for growth rates were 3–5 times lower than concentrations needed to produce maximum biomass. Modified Monod models using both DIN and SRP were developed to explain algal growth rates and peak biomass, which respectively explained 44 and 70% of the variance in algal response.     

Do secondary compounds inhibit microbial- and insect-mediated leaf breakdown in a tropical rainforest stream, Costa Rica?

We examined the hypothesis that high concentrations of secondary compounds in leaf litter of some tropical riparian tree species decrease leaf breakdown by inhibiting microbial and insect colonization. We measured leaf breakdown rates, chemical changes, bacterial, fungal, and insect biomass on litterbags of eight species of common riparian trees incubated in a lowland stream in Costa Rica. The eight species spanned a wide range of litter quality due to varying concentrations of nutrients, structural and secondary compounds. Leaf breakdown rates were fast, ranging from 0.198 d⁻¹ (Trema integerrima) to 0.011 d⁻¹ (Zygia longifolia). Processing of individual chemical constituents was also rapid: cellulose was processed threefold faster and hemicellulose was processed fourfold faster compared to similar studies in temperate streams. Leaf toughness (r = −0.86, P = 0.01) and cellulose (r = −0.78, P = 0.02) were the physicochemical parameters most strongly correlated with breakdown rate. Contrary to our initial hypothesis, secondary compounds were rapidly leached (threefold faster than in temperate studies), with all species losing all secondary compounds within the first week of incubation. Cellulose was more important than secondary compounds in inhibiting breakdown. Levels of fungal and bacterial biomass were strongly correlated with breakdown rate (fungi r = 0.64, P = 0.05; bacteria r = 0.93, P < 0.001) and changes in structural compounds (lignin r = −0.55, P = 0.01). Collector−gatherers were the dominant functional group of insects colonizing litterbags, in contrast to temperate studies where insect shredders dominate. Insect biomass was negatively correlated with breakdown rate (r = −0.70, P = 0.02), suggesting that insects did not play an important role in breakdown. Despite a wide range of initial concentrations of secondary compounds among the eight species used, we found that secondary compounds were rapidly leached and were less important than structural compounds in determining breakdown rates. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The influence of dissolved nutrients and particulate organic matter quality on microbial respiration and biomass in a forest stream

1. Although dissolved nutrients and the quality of particulate organic matter (POM) influence microbial processes in aquatic systems, these factors have rarely been considered simultaneously. We manipulated dissolved nutrient concentrations and POM type in three contiguous reaches (reference, nitrogen, nitrogen + phosphorus) of a low nutrient, third‐order stream at Hubbard Brook Experimental Forest (U.S.A). In each reach we placed species of leaves (mean C : N of 68 and C : P of 2284) and wood (mean C : N of 721 and C : P of 60 654) that differed in elemental composition. We measured the respiration and biomass of microbes associated with this POM before and after nutrient addition. 2. Before nutrient addition, microbial respiration rates and biomass were higher for leaves than for wood. Respiration rates of microbes associated with wood showed a larger response to increased dissolved nutrient concentrations than respiration rates of microbes associated with leaves, suggesting that the response of microbes to increased dissolved nutrients was influenced by the quality of their substrate. 3. Overall, dissolved nutrients had strong positive effects on microbial respiration and fungal, but not bacterial, biomass, indicating that microbial respiration and fungi were nutrient limited. The concentration of nitrate in the enriched reaches was within the range of natural variation in forest streams, suggesting that natural variation in nitrate among forest streams influences carbon mineralisation and fungal biomass.     

Effect of nutrient loading on biogeochemical and microbial processes in a New England salt marsh

Coastal marshes represent an important transitional zone between uplands and estuaries. One important function of marshes is to assimilate nutrient inputs from uplands, thus providing a buffer for anthropogenic nutrient loads. We examined the effects of nitrogen (N) and phosphorus (P) fertilization on biogeochemical and microbial processes during the summer growing season in a Spartina patens (Aiton (Muhl.)) marsh in the Narragansett Bay National Estuarine Research Reserve on Prudence Island (RI). Quadruplicate 1 m² plots were fertilized with N and P additions, N-only, P-only, or no additions. N-only addition significantly stimulated bacterial production and increased pore water NH₄⁺ and NO₃⁻ concentrations. Denitrification rates ranged from 0 to 8 mmol m⁻² day⁻¹. Fertilization had no apparent effect on soil oxygen consumption or denitrification measured in the summer in intact cores due to high core-to-core variation. P fertilization led to increased pore water dissolved inorganic phosphorus (DIP) concentrations and increased DIP release from soils. In contrast the control and N-only treatments had significant DIP uptake across the soil-water interface. The results suggest that in the summer fertilization has no apparent effect on denitrification rates, stimulates bacterial productivity, enhances pore water nutrient concentrations and alters some nutrient fluxes across the marsh surface.     

The Estimated Impact of Fungi on Nutrient Dynamics During Decomposition of Phragmites australis Leaf Sheaths and Stems

Decomposition of culms (sheaths and stems) of the emergent macrophyte Phragmites australis (common reed) was followed for 16 months in the litter layer of a brackish tidal marsh along the river Scheldt (the Netherlands). Stems and leaf sheaths were separately analyzed for mass loss, litter-associated fungal biomass (ergosterol), nutrient (N and P), and cell wall polymer concentrations (cellulose and lignin). The role of fungal biomass in litter nutrient dynamics was evaluated by estimating nutrient incorporation within the living fungal mass. After 1 year of standing stem decay, substantial fungal colonization was found. This corresponded to an overall fungal biomass of 49 ± 8.7 mg g⁻¹ dry mass. A vertical pattern of fungal colonization on stems in the canopy is suggested. The litter bag experiment showed that mass loss of stems was negligible during the first 6 months, whereas leaf sheaths lost almost 50% of their initial mass during that time. Exponential breakdown rates were −0.0039 ± 0.0004 and −0.0026 ± 0.0003 day⁻¹ for leaf sheaths and stems, respectively (excluding the initial lag period). In contrast to the stem tissue—which had no fungal colonization—leaf sheaths were heavily colonized by fungi (93 ± 10 mg fungal biomass g⁻¹ dry mass) prior to placement in the litter layer. Once being on the sediment surface, 30% of leaf sheath's associated fungal biomass was lost, but ergosterol concentrations recovered the following months. In the stems, fungal biomass increased steadily after an initial lag period to reach a maximal biomass of about 120 mg fungal biomass g⁻¹ dry mass for both plant parts at the end of the experiment. Fungal colonizers are considered to contain an important fraction of nutrients within the decaying plant matter. Fungal N incorporation was estimated to be 64 ± 13 and 102 ± 15% of total available N pool during decomposition for leaf sheaths and stems, respectively. Fungal P incorporation was estimated to be 37 ± 9 and 52 ± 15% of total available P during decomposition for leaf sheaths and stems, respectively. Furthermore, within the stem tissue, fungi are suggested to be active immobilizers of nutrients from the external environment because fungi were often estimated to contain more than 100% of the original nutrient stock.     

Comparing Effects of Nutrients on Algal Biomass in Streams in Two Regions with Different Disturbance Regimes and with Applications for Developing Nutrient Criteria

Responses of stream algal biomass to nutrient enrichment were studied in two regions where differences in hydrologic variability cause great differences in herbivory. Around northwestern Kentucky (KY) hydrologic variability constrains invertebrate biomass and their effects on algae, but hydrologic stability in Michigan (MI) streams permits accrual of high herbivore densities and herbivory of benthic algae. Multiple indicators of algal biomass and nutrient availability were measured in 104 streams with repeated sampling at each site over a 2−month period. Many measures of algal biomass and nutrient availability were positively correlated in both regions, however the amount of variation explained varied with measures of biomass and nutrient concentration and with region. Indicators of diatom biomass were higher in KY than MI, but were not related to nutrient concentrations in either region. Chl a and % area of substratum covered by Cladophora were positively correlated to nutrient concentrations in both regions. Cladophora responded significantly more to nutrients in MI than KY. Total phosphorus (TP) and total nitrogen (TN) explained similar amounts of variation in algal biomass, and not significantly more variation in biomass than dissolved nutrient concentrations. Low N:P ratios in the benthic algae indicated N as well as P may be limiting their accrual. Most observed responses in benthic algal biomass occurred in nutrient concentrations between 10 and 30 μg TP l⁻¹ and between 400 and 1000 μg TN l⁻¹.     

Nitrogen and phosphorus economy of the riparian shrub <Emphasis Type="Italic">Salix gracilistyla</Emphasis> in western Japan

Salix gracilistyla is one of the dominant plants in the riparian vegetation of the upper-middle reaches of rivers in western Japan. This species colonizes mainly sandy habitats, where soil nutrient levels are low, but shows high potential for production. We hypothesized that S.␣gracilistyla uses nutrients conservatively within stands, showing a high resorption efficiency during leaf senescence. To test this hypothesis, we examined seasonal changes in nitrogen (N) and phosphorus (P) concentrations in aboveground organs of S. gracilistyla stands on a fluvial bar in the Ohtagawa River, western Japan. The concentrations in leaves decreased from April to May as leaves expanded. Thereafter, the concentrations showed little fluctuation until September. They declined considerably in autumn, possibly owing to nutrient resorption. We converted the nutrient concentrations in each organ to nutrient amounts per stand area on the basis of the biomass of each organ. The resorption efficiency of N and P in leaves during senescence were estimated to be 44 and 46%, respectively. Annual net increments of N and P in aboveground organs, calculated by adding the amounts in inflorescences and leaf litter to the annual increments in perennial organs, were estimated to be 9.9 g and 0.83 g m⁻² year⁻¹, respectively. The amounts released in leaf litter were 6.7 g N and 0.44 g P m⁻². These values are comparable to or larger than those of other deciduous trees. We conclude that S. gracilistyla stands acquire large amounts of nutrients and release a large proportion in leaf litter.     

The influence of water chemistry on the enzymatic degradation of leaves in streams

1. Aquatic hyphomycetes degrade leaf litter in both softwater and hardwater streams. During growth on leaves, these fungi secrete an array of extracellular polysaccharidases that are differentially affected by pH. Hydrolytic enzymes exhibit acidic pH optima, whereas pectin lyases have neutral to alkaline pH optima. 2. Enzyme activities associated with microbial communities colonizing yellow poplar (Liriodendron tulipifera) leaves submerged in an acidic (pH 6.3), softwater stream were compared with those occurring in an alkaline (pH 8.2), hardwater stream. In addition to pH differences, the hardwater stream had higher nutrient concentrations and higher temperatures than the softwater stream. Conditions in the hardwater stream favoured greater microbial growth, fungal activity, rates of leaf breakdown and softening. However, activities of hydrolytic enzymes (xylanase, endocellulase, galacturonanase) were lower in the hardwater stream than in the softwater stream. Consequently, activities of these hydrolytic enzymes were not good indicators of leaf breakdown in these streams. 3. In contrast, the activities of pectin lyase were higher in the hardwater stream than in the softwater stream, corresponding to the greater rates of leaf breakdown and softening that occurred in the hardwater stream. These results support previous findings that pectin lyase is closely associated with the softening and maceration of leaf detritus and suggest that pectin degradation is a key process in the initial stages of leaf breakdown.     

Stimulation of leaf litter decomposition and associated fungi and invertebrates by moderate eutrophication: implications for stream assessment

1. We investigated the effect of moderate eutrophication on leaf litter decomposition and associated invertebrates in five reference and five eutrophied streams in central Portugal. Fungal parameters and litter N and P dynamics were followed in one pair of streams. Benthic invertebrate parameters that are considered useful in bioassessment were estimated in all streams. Finally, we evaluated the utility of decomposition as a tool to assess stream ecosystem functional integrity. 2. Decomposition of alder and oak leaves in coarse mesh bags was on average 2.3–2.7× faster in eutrophied than in reference streams. This was attributed to stimulation of fungal activity (fungal biomass accrual and sporulation of aquatic hyphomycetes) by dissolved nutrients. These effects were more pronounced for oak litter (lower quality substrate) than alder. N content of leaf litter did not differ between stream types, while P accrual was higher in the eutrophied than in the reference stream. Total invertebrate abundances and richness associated with oak litter, but not with alder, were higher in eutrophied streams. 3. We found only positive correlations between stream nutrients (DIN and SRP) and leaf litter decomposition rates in both fine and coarse mesh bags, associated sporulation rates of aquatic hyphomycetes and, in some cases, total invertebrate abundances and richness. 4. Some metrics based on benthic invertebrate community data (e.g. % shredders, % shredder taxa) were significantly lower in eutrophied than in reference streams, whereas the IBMWP index that is specifically designed for the Iberian peninsula classified all 10 streams in the highest possible class as having ‘very good’ ecological conditions. 5. Leaf litter decomposition was sufficiently sensitive to respond to low levels of eutrophication and could be a useful functional measure to complement assessment programmes based on structural parameters.     

Litter decomposition in a Cerrado savannah stream is retarded by leaf toughness, low dissolved nutrients and a low density of shredders

1. To assess whether the reported slow breakdown of litter in tropical Cerrado streams is due to local environmental conditions or to the intrinsic leaf characteristics of local plant species, we compared the breakdown of leaves from Protium brasiliense, a riparian species of Cerrado (Brazilian savannah), in a local and a temperate stream. The experiment was carried out at the time of the highest litter fall in the two locations. An additional summer experiment was conducted in the temperate stream to provide for similar temperature conditions. 2. The breakdown rates (k) of P. brasiliense leaves in the tropical Cerrado stream ranged from 0.0001 to 0.0008 day⁻¹ and are among the slowest reported. They were significantly (F = 20.12, P < 0.05) lower than in the temperate stream (0.0046–0.0055). The maximum ergosterol content in decomposing leaves in the tropical Cerrado stream was 106 μg g⁻¹, (1.9% of leaf mass) measured by day 75, which was lower than in the temperate stream where maximum ergosterol content of 522 μg g⁻¹ (9.5% of leaf mass) was achieved by day 30. The ATP content, as an indicator of total microbial biomass, was up to four times higher in the tropical Cerrado than in the temperate stream (194.0 versus 49.4 nmoles g⁻¹). 3. Unlike in the temperate stream, leaves in the tropical Cerrado were not colonised by shredder invertebrates. However, in none of the experiments did leaves exposed (coarse mesh bags) and unexposed (fine mesh bags) to invertebrates differ in breakdown rates (F = 1.15, P > 0.05), indicating that invertebrates were unable to feed on decomposing P. brasiliense leaves. 4. We conclude that the slow breakdown of P. brasiliense leaves in the tropical Cerrado stream was because of the low nutrient content in the water, particularly nitrate (0.05 mgN L⁻¹), which slows down fungal activity and to the low density of invertebrates capable of using these hard leaves as an energy source.     

Interaction Effects of Ambient UV Radiation and Nutrient Limitation on the Toxic Cyanobacterium <Emphasis Type="Italic">Nodularia spumigena</Emphasis>

Nodularia spumigena is one of the dominating species during the extensive cyanobacterial blooms in the Baltic Sea. The blooms coincide with strong light, stable stratification, low ratios of dissolved inorganic nitrogen, and dissolved inorganic phosphorus. The ability of nitrogen fixation, a high tolerance to phosphorus starvation, and different photo-protective strategies (production of mycosporine-like amino acids, MAAs) may give N. spumigena a competitive advantage over other phytoplankton during the blooms. To elucidate the interactive effects of ambient UV radiation and nutrient limitation on the performance of N. spumigena, an outdoor experiment was designed. Two radiation treatments photosynthetic active radiation (PAR) and PAR +UV-A + UV-B (PAB) and three nutrient treatments were established: nutrient replete (NP), nitrogen limited (−N), and phosphorus limited (−P). Variables measured were specific growth rate, heterocyst frequency, cell volume, cell concentrations of MAAs, photosynthetic pigments, particulate carbon (POC), particulate nitrogen (PON), and particulate phosphorus (POP). Ratios of particulate organic matter were calculated: POC/PON, POC/POP, and PON/POP. There was no interactive effect between radiation and nutrient limitation on the specific growth rate of N. spumigena, but there was an overall effect of phosphorus limitation on the variables measured. Interaction effects were observed for some variables; cell size (larger cells in −P PAB compared to other treatments) and the carotenoid canthaxanthin (highest concentration in −N PAR). In addition, significantly less POC and PON (mol cell⁻¹) were found in −P PAR compared to −P PAB, and the opposite radiation effect was observed in −N. Our study shows that despite interactive effects on some of the variables studied, N. spumigena tolerate high ambient UVR also under nutrient limiting conditions and maintain positive growth rate even under severe phosphorus limitation.     

Seasonal variation in nutrient limitation of microbial biofilms colonizing organic and inorganic substrata in streams

Humans have increased the availability of nutrients including nitrogen and phosphorus worldwide; therefore, understanding how microbes process nutrients is critical for environmental conservation. We examined nutrient limitation of biofilms colonizing inorganic (fritted glass) and organic (cellulose sponge) substrata in spring, summer, and autumn in three streams in Michigan, USA. Biofilms were enriched with nitrate (NO₃ ⁻), phosphate (PO₄ ³⁻), ammonium (NH₄ ⁺), NO₃ ⁻ + PO₄ ³⁻, NH₄ ⁺ + PO₄ ³⁻, or none (control). We quantified biofilm structure and function as chlorophyll a (i.e., primary producer biomass) and community respiration on all substrata. In one stream, we characterized bacterial and fungal communities on cellulose in autumn using clone library sequencing and denaturing gradient gel electrophoresis to determine if community structure was linked to nutrient limitation status. Despite oligotrophic conditions, primary producer biomass was infrequently nutrient limited. In contrast, respiration on organic substrata was frequently limited by N + P combinations. We found no difference between biofilm response to NH₄ ⁺ versus NO₃ ⁻ enrichment, although the response to both N-species was positively related to water column PO₄ ³⁻ concentrations and temperature. Molecular analysis for fungal community composition suggested no relationship to nutrient limitation, but the dominant members of the bacterial community on cellulose were different on NO₃ ⁻, PO₄³, and NO₃ ⁻ + PO₄ ³⁻ treatments relative to control, NH₄ ⁺, and NH₄ ⁺ + PO₄ ³⁻ treatments, which matched patterns for biofilm respiration rates from each treatment. Our results show discrete patterns of nutrient limitation dependent upon substratum type and season, and imply changes in bacterial community structure and function may be linked following nutrient enrichment in streams.     

Leaf litter breakdown budgets in streams of various trophic status: effects of dissolved inorganic nutrients on microorganisms and invertebrates

1. We investigated the effect of trophic status on the organic matter budget in freshwater ecosystems. During leaf litter breakdown, the relative contribution of the functional groups and the quantity/quality of organic matter available to higher trophic levels are expected to be modified by the anthropogenic release of nutrients. 2. Carbon budgets were established during the breakdown of alder leaves enclosed in coarse mesh bags and submerged in six streams: two oligotrophic, one mesotrophic, two eutrophic and one hypertrophic streams. Nitrate concentrations were 4.5–6.7 mg L⁻¹ and the trophic status of each stream was defined by the soluble reactive phosphorus concentration ranging from 3.4 (oligotrophic) to 89 μg L⁻¹ (hypertrophic). An ammonium gradient paralleled the phosphate gradient with mean concentrations ranging from 1.4 to 560 μg L⁻¹ NH₄‐N. The corresponding unionised ammonia concentrations ranged from 0.08 to 19 μg L⁻¹ NH₃‐N over the six streams. 3. The dominant shredder taxa were different in the oligo‐, meso‐ and eutrophic streams. No shredders were observed in the hypertrophic stream. These changes may be accounted for by the gradual increase in the concentration of ammonia over the six streams. The shredder biomass dramatically decreased in eu‐ and hypertrophic streams compared with oligo‐ and mesotrophic. 4. Fungal biomass increased threefold from the most oligotrophic to the less eutrophic stream and decreased in the most eutrophic and the hypertrophic. Bacterial biomass increased twofold from the most oligotrophic to the hypertrophic stream. Along the trophic gradient, the microbial CO₂ production followed that of microbial biomass whereas the microbial fine particulate organic matter and net dissolved organic carbon (DOC) did not consistently vary. These results indicate that the microorganisms utilised the substrate and the DOC differently in streams of various trophic statuses. 5. In streams receiving various anthropogenic inputs, the relative contribution of the functional groups to leaf mass loss varied extensively as a result of stimulation and the deleterious effects of dissolved inorganic compounds. The quality/quantity of the organic matter produced by microorganisms slightly varied, as they use DOC from stream water instead of the substrate they decompose in streams of higher trophic status.     

Nutrients and temperature additively increase stream microbial respiration

Rising temperatures and nutrient enrichment are co‐occurring global‐change drivers that stimulate microbial respiration of detrital carbon, but nutrient effects on the temperature dependence of respiration in aquatic ecosystems remain uncertain. We measured respiration rates associated with leaf litter, wood, and fine benthic organic matter (FBOM) across seasonal temperature gradients before (PRE) and after (ENR1, ENR2) experimental nutrient (nitrogen [N] and phosphorus [P]) additions to five forest streams. Nitrogen and phosphorus were added at different N:P ratios using increasing concentrations of N (~80–650 μg/L) and corresponding decreasing concentrations of P (~90–11 μg/L). We assessed the temperature dependence, and microbial (i.e., fungal) drivers of detrital mass‐specific respiration rates using the metabolic theory of ecology, before vs. after nutrient enrichment, and across N and P concentrations. Detrital mass‐specific respiration rates increased with temperature, exhibiting comparable activation energies (E, electronvolts [eV]) for all substrates (FBOM E = 0.43 [95% CI = 0.18–0.69] eV, leaf litter E = 0.30 [95% CI = 0.072–0.54] eV, wood E = 0.41 [95% CI = 0.18–0.64] eV) close to predicted MTE values. There was evidence that temperature‐driven increased respiration occurred via increased fungal biomass (wood) or increased fungal biomass‐specific respiration (leaf litter). Respiration rates increased under nutrient‐enriched conditions on leaves (1.32×) and wood (1.38×), but not FBOM. Respiration rates responded weakly to gradients in N or P concentrations, except for positive effects of P on wood respiration. The temperature dependence of respiration was comparable among years and across N or P concentration for all substrates. Responses of leaf litter and wood respiration to temperature and the combined effects of N and P were similar in magnitude. Our data suggest that the temperature dependence of stream microbial respiration is unchanged by nutrient enrichment, and that increased temperature and N + P availability have additive and comparable effects on microbial respiration rates.     

Biomass production and nutrient accumulation in <Emphasis Type="Italic">Sparganium emersum</Emphasis> Rehm. after sediment treatment with mineral and organic fertilisers in three mesocosm experiments

The response of the aquatic plant Sparganium emersum to different sediment nutrient levels was studied in three mesocosm experiments. The aim was to assess plant growth parameters and nutrient accumulation in the plant tissue under conditions relevant for habitats with sediments affected by anthropogenic nutrient enrichment. The experimental treatments were produced by fertilisation of the rooting medium (washed river sand) with differing doses of either NPK mineral fertiliser or digested sludge from solid pig slurry waste. Growth inhibition by high nutrient levels was not observed in any treatment (highest nutrient concentrations in the sediment with mineral fertiliser: N 250 mg kg⁻¹, P 50 mg kg⁻¹; organic fertiliser: N 6300 mg kg⁻¹, P 1800 mg kg⁻¹), which confirms the tolerance of S. emersum to high nutrient loads. The sediment nutrient concentration was best reflected in shoot dry mass. Nutrient contents in plant tissues were similar for most nutrient concentrations in the rooting media; only N increased significantly with N levels in the sediment in belowground parts. Nutrient standing stocks in plants, however, generally corresponded to the nutrient supply, and reached highest values (max. N 3.7 g m⁻², P 1.2 g m⁻²) in the richest treatments with organic fertiliser. The capability of S. emersum to use nutrients from high sediment concentrations and in organically polluted environments recommends this species for use in water quality management including tertiary wastewater treatment.