Microfaunal response to fertilization of an arctic tundra stream

1. As part of a whole-system study, the response of the heterotrophic microfaunal community colonizing artificial substrata (polyfoam units) to fertilization of an arctic tundra stream was followed for 6 weeks during the summer. 2. Dominant heterotrophic microfauna observed included amoebae (approximately 40% of colonizing biomass), rotifers (36% of biomass) and ciliates (25% of biomass). 3. Biomass of heterotrophic microfauna on artificial substrata was not significantly different in a control reach and an experimental reach fertilized with phosphorus (loading rate ten times ambient), but in a reach fertilized with both phosphorus and nitrogen (loading rates ten times ambient) biomass was double that of the control and phosphorus-fertilized reaches. The lack of response in the phosphorus reach was probably due to greater insect grazing as a result of previous phosphorus fertilization of this reach. 4. Abundance of microfauna on epilithic surfaces in the river was higher on rocks from pools than on rocks from riffle areas, but abundance on the artificial substrata was higher than on the natural rocks. 5. The results suggest that microfauna of arctic tundra streams are regulated by grazers and that their importance in transfers among trophic levels is greater in pools than in riffles.     

Recovery of three arctic stream reaches from experimental nutrient enrichment

1. Nutrient enrichment and resulting eutrophication is a widespread anthropogenic influence on freshwater ecosystems, but recovery from nutrient enrichment is poorly understood, especially in stream environments. We examined multi‐year patterns in community recovery from experimental low‐concentration nutrient enrichment (N + P or P only) in three reaches of two Arctic tundra streams (Kuparuk River and Oksrukuyik Creek) on the North Slope of Alaska (U.S.A.). 2. Rates of recovery varied among community components and depended on duration of enrichment (2–13 consecutive growing seasons). Biomass of epilithic algae returned to reference levels rapidly (within 2 years), regardless of nutrients added or enrichment duration. Aquatic bryophyte cover, which increased greatly in the Kuparuk River only after long‐term enrichment (8 years), took 8 years of recovery to approach reference levels, after storms had scoured most remnant moss in the recovering reach. 3. Multi‐year persistence of bryophytes in the Kuparuk River appeared to prevent recovery of insect populations that had either been positively (e.g. the mayfly Ephemerella, most chironomid midge taxa) or negatively (e.g. the tube‐building chironomid Orthocladius rivulorum) affected by this shift in dominant primary producer. These lags in recovery (of >3 years) were probably driven by the persistent effect of bryophytes on physical benthic habitat. 4. Summer growth rates of Arctic grayling (both adults and young‐of‐year) in Oksrukuyik Creek (fertilised for 6 years with no bryophyte colonisation), which were consistently increased by nutrient addition, returned to reference rates within 1–2 years. 5. Rates of recovery of these virtually pristine Arctic stream ecosystems from low‐level nutrient enrichment appeared to be controlled largely by duration of enrichment, mediated through physical habitat shifts caused by eventual bryophyte colonisation, and subsequent physical disturbance that removed bryophytes. Nutrient enrichment of oligotrophic Arctic stream ecosystems caused by climate change or local anthropogenic activity may have dramatic and persistent consequences if it results in the colonisation of long‐lived primary producers that alter physical habitat.     

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

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

Black fly (Diptera: Simuliidae) response to phosphorus enrichment of an arctic tundra stream

This study reports on the density, growth, and production response of the dominant black fly, Prosimulium martini, to whole river fertilization of the Kuparuk River in arctic Alaska during the summer of 1984. Beginning in 1983, a long term study of fertilization effects was initiated in the Kuparuk River. Increased nutrient supply stimulated algal and microbial biomass and microbial activity, which in turn affected the larval growth and abundance of Prosimulium. This experiment allowed us to isolate the effects of nutrient supply from other factors in determining black fly growth and abundance. Phosphorus addition had the following indirect effects on Prosimulium: growth was higher, but abundance decreased in the enriched section, leading to a net decrease in secondary production from 2.62 g m⁻² yr⁻¹ to 0.77 g m⁻² yr⁻¹. Prosimulium emergence rates were not measureably affected. The decrease in abundance and production appears to be a result of competitive displacement by the caddisfly Brachycentrus americanus which increased in abundance in the fertilized section of the river.     

The trophic interactions of young Arctic grayling (Thymallus arcticus) in an Arctic tundra stream

1. Young (0+) Arctic grayling (Thymallus arcticus) have the potential to control the trophic structure of Arctic tundra streams through consumption, nutrient excretion and the modification of prey behaviour. The effect of young grayling on three trophic levels (algae, invertebrates and fish) was investigated by manipulating fish density and by fertilizing the river with phosphorus (P). 2. Nutrients, epilithic chlorophyll a, benthic invertebrates and fish biomass were measured within each fish density treatment (0, 4, and 40 fish m^–2) within the P-limited reference zone and the P-enriched fertilized zone of the Kuparuk River, Alaska. 3. Epilithic chlorophyll a increased with increased fish density in both reference and fertilized zones, while mayfly density decreased with increased fish density in the fertilized zone only. Final mean mass of young grayling in the 40 fish m^–2 cages was lower than mean mass in the 4 fish m^–2 cages. 4. Young grayling may produce a top-down cascading trophic effect in areas where nutrients are not limited. 5. River nutrient status and river discharge may modify the strength of top-down control by young grayling.     

Nitrogen and phosphorus dynamics in hot desert streams of Southwestern U.S.A.

Nitrogen to phosphorus ratios and concentrations of nitrate and soluble reactive phosphate are presented for an array of Southwestern streams as evidence that nitrogen is the limiting nutrient where such limitation occurs. Nitrate uptake in sections of intermittent streams was attributable to autotrophic activity. Uptake of soluble reactive phosphate was unrelated to any indicator of autotrophic activity, thus concentrations of this nutrient in desert and semi-desert stream waters may be controlled by other factors.     

Functional responses of stream biofilms to flow cessation,desiccation and rewetting

1. Mediterranean climate regions are characterised by long summer droughts that usually involve flow intermittency in low‐ to mid‐order streams. Flow intermittency implies flow cessation, drying and subsequent rewetting of the streambed, and affects both autotrophic and heterotrophic processes. The balance between these processes, as well as the balance in the use of carbon (C), nitrogen (N) and phosphorus (P) may change because of the ongoing increase in stream flow intermittency caused by global change in many regions. It is therefore crucial to understand better the consequences of this phenomenon. 2. Our two initial hypotheses were (i) that flow intermittency would impact more on autotrophic than on heterotrophic processes in stream biofilms owing to the higher water dependence of autotrophs, as well as differences in the water storage capacity of the stream biofilm compartments where autotrophic and heterotrophic processes mainly occur (surface cobbles versus hyporheic sediments) and (ii) that the C‐N‐P use by biofilms would change during the dry period (terrestrial phase) owing to the extreme water stress conditions. These hypotheses were tested by analysing the functional response of the main stream biofilms (epilithic, epipsammic and hyporheic) during flow cessation, desiccation and rewetting in a Mediterranean forested stream. The autotrophic response was characterised through changes in the photon yield, whereas the heterotrophic response was characterised by changes in the extracellular enzyme activities. 3. Streambed desiccation had clear effects on the functioning of stream biofilms. Autotrophic biomass decreased by 80% with streambed desiccation, but recovered rapidly after flow resumption. Heterotrophs were more resistant to water stress, especially in the epipsammic and hyporheic biofilms where bacterial cell density decreased only by 20%. 4. Extracellular enzyme activities remained relatively high, and the balance in the C‐N‐P use by biofilms changed during the dry period. The C and P breakdown capacities were maintained during dry conditions, especially in the epipsammic and hyporheic biofilms, but the degradation of N compounds sharply decreased. Elemental molar ratios (C:N and C:P) of the different biofilms also changed with streambed desiccation. C:P ratios increased from 80 to 300, while the C:N ratios increased from 10 to 16. 5. Given the contrasting responses of autotrophic and heterotrophic processes in the different biofilms, our results suggest that the current increase in flow intermittency extent is likely to increase the relative importance of heterotrophic processes in stream ecosystems, as well as the relative contribution of the hyporheic biofilm to C‐N‐P use. Our results further suggest that the longer streams remain dry, the more the biofilm stoichiometry will change.     

Seasonal changes in composition and biomass of epilithic algal floras of a chalk stream and a soft water stream with estimates of production

Microscopic epilithic algae in the River Itchen at Otterbourne near Southampton and in the Ober Water in the New Forest were studied during 1984 and 1985. The River Itchen rises from chalk springs and has a steady pH near 8.2 and a mean alkalinity of 236 mg HCO₃ 1^–1; at the study site the river is about 16 m wide and 20 cm deep, with a mean flow rate of 0.33 m s^–1 and a discharge ranging through the year between 0.34 and 2.46 m³ s^–1. The Ober Water, which drains sands and gravels, has a pH between 6.9 and 7.2 and a mean alkalinity of about 50 mg HCO₃ 1^–1; at the study site it is about 6 m wide, with a mean flow rate of 0.27 m s^–1 and a discharge ranging through the year between 0.08 and 1.0 m³ s^–1.Epilithic algae removed from the pebbles that form the major part of the beds of both streams show seasonal changes in abundance and composition. Diatoms peaked in April/May and dominate the epilithic flora in both streams, comprising 70–95% of all algal cells; highest numbers of chlorophytes occurred in summer and cyanophytes increased in autumn. The species composition of the epilithic flora in the two streams was different, as was the population density; algal cell numbers ranged between 500 and 7000 cells mm^–2 of stream floor in the River Itchen and between 8 and 320 cells mm^–2 of stream floor in the Ober Water. The chlorophyll a content of epilithic algae in the River Itchen ranged between 115 and 415 mg m^–2 of stream floor, representing an annual mean biomass of about 8 g m^–2, whereas in the Ober Water a chlorophyll a content of 2.2 to 44 mg m^–2 of stream floor was found, representing an annual mean biomass of about 1 g m^–2. Cautious estimates of the annual production of epilithic algae in these streams suggest a value of about 600 g organic dry weight m^–2 in the River Itchen and about 75 g m^–2 in the Ober Water.     

Does nitrogen become limiting under high‐P conditions in detritus‐based tropical streams?

1. We examined effects of nutrients on leaf breakdown in interior forest streams at La Selva Biological Station, Costa Rica. We tested the hypothesis that dissolved inorganic nitrogen (DIN) becomes limiting when ambient phosphorus (P) concentration is high. We also compared the breakdown of relatively ‘low quality’ leaves (lower C : N, Trema integerrima) with that of ‘higher quality’ leaves (higher C : N, Ficus insipida) in a high‐P stream. 2. Litterbags were incubated in two streams: one enriched experimentally with P [target concentration 200 μg soluble reactive phosphorus (SRP) L^?1] and one control (naturally low P concentration approximately 10 μg SRP L^?1). Ammonium enrichment was achieved by adding fertiliser upstream of half of the litterbags in each stream. 3. Phosphorus addition stimulated leaf breakdown, microbial respiration, ergosterol and leaf %P. Leaf breakdown rate was consistent with those in La Selva streams with naturally high P concentration. 4. Nitrogen (N) addition had no effect on leaf breakdown, microbial respiration, ergosterol or leaf chemistry in either the P‐enriched or the reference stream, in spite of low N : P ratios. We conclude that N is probably not limiting in streams at La Selva that are naturally high in P. This may be due to moderately high ambient N concentration (>200 μg DIN L^?1) prevailing throughout the year. 5. The species with a lower C : N decomposed more rapidly and supported higher microbial activity than that with a higher C : N. Subtle differences in leaf N content, as well as dissolved P concentration, may be important in determining microbial colonisation and subsequent leaf breakdown.     

Carbon, nitrogen, and phosphorus dynamics during leaf decay in nutrient-enriched stream microecosystems

We investigated the effects of long-term enrichment with nitrate, phosphate, and nitrate+phosphate on the first 5 weeks of leaf detritus processing in laboratory stream microecosystems. Enrichment with nitrate+phosphate accelerated leaf weight loss and increased rates of respiration associated with the leaves. However, whole-system respiration was little changed from that observed in the control stream since respiration in the water was greatly reduced. Enrichment with phosphate alone had little effect except to lower respiration associated with leaf discs. Enrichment with nitrate alone also decreased leaf-disc respiration but resulted in a greatly increased rate of respiration in the water. Net leaching and fragmentation of carbon from the leaves was also increased by nitrate enrichment. Nitrogen and phosphorus levels in leaf material were little affected by enrichment with nitrate or phosphorus alone. Leaves in those streams and in the control stream released nitrogen and phosphorus to the water. In contrast, percent nitrogen and phosphorus increased greatly in the leaves in the stream enriched with both nitrate and phosphate. The leaves in this system immobilized both nitrogen and phosphorus from the water. We also studied the importance of nitrogen fixation as a vector for nitrogen incorporation associated with leaf decomposition in streams. Somewhat surprisingly, fixation by microbes associated with the leaves and by microbes suspended in the water occurred under all three experimental enrichment treatments as well as in the control, casting doubt on the effectiveness of nitrate in inhibiting nitrogenase synthesis in nature. However, N₂-fixation is only a minor source of nitrogen for leaves decaying under the conditions studied.     

A stressed stream ecosystem: macroinvertebrate community integrity and microbial trophic response

A year-long study of a second-order stream in Southwestern Virginia was carried out from 1979–80. One of the objectives of the study was to evaluate the effects of sewage and electroplating plant effluent stress on the trophic response of aquatic invertebrate assemblages and microbial communities in the stream. Quantitative benthic samples were collected periodically at three reference stations and four stressed stations below the outfalls. Invertebrates were counted, identified taxonomically, and classified into functional groups based on their feeding strategies. Ash-free dry weights were obtained for each functional group by date and station, and the number and density of different taxa were calculated as well. Reference stations had diverse invertebrate assemblages; scrapers were well represented and all functional groups were present in reasonably equivalent proportions. Stressed stations were dominated by collector gatherers and filterers to the virtual exclusion of scrapers. The trophic status of the microbial community was determined by suspending artificial substrates in the stream for 1-week periods. The community that colonized the substrates was assayed for ATP and chlorophyll a, and an autotrophy index (AI) was calculated using these values. The autotrophic component of the microbial community was greatest at the reference stations, and the community became primarily heterotrophic below the outfalls. The AI correlated well with the proportion of scrapers. Aquatic invertebrate assemblages and microbial communities responded to stress by changing their trophic structure to fit best the available energy sources. Where heterotrophic microbes dominated, gathering and filtering invertebrates utilized the abundant organic matter. In areas where a mainly autotrophic microbial community existed, scrapers, gatherers, and filterers were all present in balanced proportions.     

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.     

Seasonal relationships between planktonic microorganisms and dissolved organic material in an alpine stream

The relationships between the abundance and activity of planktonic, heterotrophic microorganisms and the quantity and characteristics of dissolved organic carbon (DOC) in a Rocky Mountain stream were evaluated. Peak values of glucose uptake, 2.1 nmol L⁻¹ hr⁻¹, and glucose concentration, 333 nM, occurred during spring snowmelt when the water temperature was 4.0°C and the DOC concentration was greatest. The turnover time of thein situ glucose pool ranged seasonally from 40–1110 hours, with a mean of 272 hr. Seasonal uptake of³H-glucose, particulate ATP concentrations, and direct counts of microbial biomass were independent of temperature, but were positively correlated with DOC concentrations and negatively correlated with stream discharge. Heterotrophic activity in melted snow was generally low, but patchy. In the summer, planktonic heterotrophic activity and microbial biomass exhibited small-scale diel cycles which did not appear to be related to fluctuations in discharge or DOC, but could be related to the activity of benthic invertebrates. Leaf-packs placed under the snow progressively lost weight and leachable organic material during the winter, indicating that the annual litterfall in the watershed may be one source of the spring flush of DOC. These results indicate that the availability of labile DOC to the stream ecosystem is the primary control on seasonal variation in heterotrophic activity of planktonic microbial populations.     

Nitrogen limitation of heterotrophic biofilms in boreal streams

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Nitrogen uptake and transformation in a midwestern U.S. stream: A stable isotope enrichment study

This study presents a comprehensive analysis ofnitrogen (N) cycling in a second-order forestedstream in southern Michigan that has moderatelyhigh concentrations of ammonium (mean,16 g N/L) and nitrate (17 g N/L). Awhole-stream ¹⁵NH₄ ⁺ addition wasperformed for 6 weeks in June and July, and thetracer ¹⁵N was measured downstream inammonium, nitrate, and detrital and livingbiomass. Ancillary measurements includedbiomass of organic matter, algae, bacteria andfungi, nutrient concentrations, hydrauliccharacteristics, whole-stream metabolism, andnutrient limitation assays. The resultsprovide insights into the heterotrophic natureof woodland streams and reveal the rates atwhich biological processes alter nitrogentransport through stream systems.Ammonium uptake lengths were 766–1349 m anduptake rates were 41–60 g N m^–2min^–1. Nitrate uptake could not bedetected. Nitrification rates were estimatedfrom the downstream increase in¹⁵N-enriched nitrate using a simulationmodel. The ammonium was removed bynitrification (57% of total uptake),heterotrophic bacteria and fungi associatedwith detritus (29%), and epilithic algae(14%). Growth of algae was likely limited bylight rather than nutrients, and dissolvedO₂ revealed that the stream metabolism washeterotrophic overall (P:R = 0.2). Incubationsof detritus in darkened chambers showed thatuptake of ¹⁵N was mostly heterotrophic.Microbial N in detritus and algal N inepilithon appeared to reach isotopic steadystate with the dissolved ammonium, but theisotopic enrichment of the bulk detritus andepilithon did not approach that of ammonium,probably due to a large fraction of organic Nin the bulk samples that was not turning over. The actively cycling fraction of total N inorganic compartments was estimated from theisotopic enrichment, assuming uptake ofammonium but not nitrate, to be 23% forepilithon, 1% for fine benthic organic matter,5% for small woody debris, and 7% for leaves. These percentages agree with independentestimates of epilithic algal biomass, whichwere based on carbon:chlorophyll ratios in bulksamples and in algal fractions separated bydensity-gradient centrifugation in colloidalsilica, and of microbial N in the detritus,which were based on N released by chloroformfumigations.     

The influence of elevated nitrate concentration on rate of leaf decomposition in a stream

SUMMARY.

• 1 Leaf decomposition was compared in two streams at the Coweeta Hydrologic Laboratory, North Carolina. U.S.A. One stream drains an undisturbed hardwood watershed, while the other drains a successional watershed subject to an insect outbreak. The successional watershed has elevated nitrate concentrations in the streamwater.
• 2 Both black locust (Robinia pseudo-acacia) and sweet birch (Betula lenta) leaf litter decomposed 2.8 times more rapidly in the stream with high nitrate concentrations.
• 3 The more rapid decay rates appeared to be partly due to accelerated microbial processing in response to nitrate enrichment, because microbial biomass (as ATP) was higher in the nitrate-enriched stream.
• 4 At each point in time, nitrogen and phosphorus content of the litter was lower in the high nitrate stream; however, there was no significant difference in nitrogen or phosphorus content at the same state of leaf decay in the two streams.

     

Leaf litter decomposition and microbial activity in nutrient-enriched and unaltered reaches of a headwater stream

SUMMARY 1. Decomposition of red maple ( Acer rubrum ) and rhododendron ( Rhododendron maximum ) leaves and activity of associated microorganisms were compared in two reaches of a headwater stream in Coweeta Hydrologic Laboratory, NC, U.S.A. The downstream reach was enriched with ammonium, nitrate, and phosphate whereas the upstream reach was not altered.
2. Decomposition rate, microbial respiration, fungal and bacterial biomass, and the sporulation rate of aquatic hyphomycetes associated with decomposing leaf material were significantly higher for both leaf types in the nutrient-enriched reach. Species richness and community structure of aquatic hyphomycetes also exhibited considerable changes with an increase in the number of fungal codominants in the nutrient-enriched reach.
3. Fungal biomass was one to two orders of magnitude greater than bacterial biomass in both reaches. Changes in microbial respiration rate corresponded to those in fungal biomass and sporulation, suggesting a primary role of fungi in leaf decomposition.
4. Nutrient enrichment increased microbial activity, the proportion of leaf carbon channelled through the microbial compartment and the decomposition rate of leaf litter.     

Meiofaunal responses to nutrient additions in a Mediterranean stream

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

Effects of phosphorus enrichment and grazing snails on modern stromatolitic microbial communities

1. The effects of phosphorus enrichment and grazing snails on a benthic microbial community that builds stromatolic oncolites were examined in an experiment at Rio Mesquites, Cuatro Ciénegas, Mexico. Chemical analyses of stream water samples indicated that overall atomic ratios of total nitrogen (N) to total phosphorus (P) were approximately 110, indicating a strong potential for P‐limitation of microbial growth. 2. Phosphorus enrichment involved addition of 5 μmol Na₂HPO₄ L^?1 to streamside microcosms receiving intermittent inputs of stream water while grazer manipulation involved removal of the dominant grazer, the snail Mexithauma quadripaludium. After 7 weeks, we examined responses in organic matter content, C : N : P ratios, metabolism (P removal, primary production, dark respiration, and calcification), and microbial community structure using molecular fingerprinting of 16S rRNA genes. 3. Manipulation of snails did not affect response variables measured in these treatments (organic matter, C : P ratio, P removal rate). However, P enrichment significantly decreased the C : P and N : P ratios of surficial materials in the oncolites (organic matter content was unchanged), increased net and gross photosynthesis (oxygen consumption in the dark was unchanged), increased rates of calcification, and increased diatoms relative to cyanobacteria. Heterotrophic Eubacteria and Archaea were only modestly affected. Thus, our results indicate weak grazing effects but strong impacts of P in this benthic system. 4. We hypothesise that a state of severe P‐limitation is imposed on autotrophic production in this food web due, at least in part, to co‐precipitation of phosphate during calcite deposition. This produces severe P‐limitation of the benthic algae and cyanobacteria, resulting in high C : P ratio of microbial mats relative to the biomass of photoautotrophs (phytoplankton, terrestrial foliage) in other ecosystems. In turn, this high C : P ratio is likely to generate severe stoichiometric constraints on the herbivores, thus limiting their populations and resulting in weak overall grazing impacts.     

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

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