Continuous ammonium enrichment of a woodland stream: uptake kinetics, leaf decomposition, and nitrification

• 1 In order to test for nitrogen limitation and examine ammonium uptake by stream sediments, ammonium hydroxide was added continuously at concentrations averaging 100 μg1^-1 for 70 days to a second- order reach of Walker Branch, an undisturbed woodland stream in Tennessee.
• 2 Ammonium uptake during the first 4h of addition corresponded to adsorption kinetics rather than to first-order uptake or to Michaelis- Menten kinetics. However, the calculated adsorption partition coefficient was two to four orders of magnitude greater than values reported for physical adsorption of ammonium, suggesting that the uptake was largely biotic.
• 3 Mass balance indicated that the uptake of ammonium from the water could be accounted for by increased nitrogen content in benthic organic detritus. Nitrification, inferred from longitudinal gradients in NO₃, began soon after enrichment and increased dramatically near the end of the experiment.
• 4 Both ammonium and nitrate concentrations dropped quickly to near background levels when input ceased, indicating little desorption or nitrification of excess nitrogen stored in the reach.
• 5 There was no evidence of nitrogen limitation as measured by weight loss, oxygen consumption, phosphorus content, and macroinvertebrate density of red oak leaf packs, or by chlorophyll content and aufwuchs biomass on plexiglass slides. A continuous phosphorus enrichment 1 year earlier had demonstrated phosphorus limitation in Walker Branch.

     

Seston quality drives feeding,stoichiometry and excretion of zebra mussels

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Recruitment preferences of blue mussel spat (Mytilus edulis) for different substrata and microhabitats in the White Sea (Russia)

We adapted the chloroform fumigation method to determine microbial nitrogen (N) and microbial incorporation of ¹⁵N on three common substrates [leaves, wood and fine benthic organic matter (FBOM)] in three forest streams. We compared microbial N and ¹⁵N content of samples collected during a 6-week ¹⁵N–NH₄ tracer addition in each stream. The ¹⁵N was added during late autumn to Upper Ball Creek, a second-order stream at the Coweeta Hydrologic Lab, North Carolina, U.S.A.; during spring to Walker Branch, a first-order stream on DOE's Oak Ridge National Environmental Research Park, Tennessee; and during summer to Bear Brook, a first-order stream in the Hubbard Brook Experimental Forest, New Hampshire. FBOM was the largest component of organic matter and N standing stock in all streams. Microbial N represented the highest proportion of total N in leaves and least in FBOM in Walker Branch and Bear Brook. In Upper Ball Creek, the proportion of microbial N was higher in FBOM than in used biofilm or on leaves. Standing stock of microbial N on leaves and in FBOM ranged from 37 mg N m^–2 in Bear Brook to 301 mg N m^–2 in Walker Branch. Percent of detrital N in living microbial cells was directly related to total microbial biomass (fungal and bacterial biomass) determined from microscopic counts. ¹⁵N values for microbes were generally higher than for bulk detritus, which would result in higher ¹⁵N values for animals preferentially consuming or assimilating microbial cells. The proportion of ¹⁵N taken up by detritus during the ¹⁵N experiments that remained in microbial cells by the end of the experiments was highest for wood biofilm in Upper Ball Creek (69%), leaves in Walker Branch (65%) and FBOM in Upper Ball Creek (31%). Lower retention proportions (<1–25%) were observed for other substrates. Our results suggest that microbial cells associated with leaves and wood biofilm were most active in ¹⁵N–NH₄ immobilization, whereas microbial cells associated with FBOM immobilized little ¹⁵N from stream water.     

Experimental nutrient enrichment of forest streams increases energy flow to predators along greener food‐web pathways

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Temporal Variation in the Importance of a Dominant Consumer to Stream Nutrient Cycling

Animal excretion can be a significant nutrient flux within ecosystems, where it supports primary production and facilitates microbial decomposition of organic matter. The effects of excretory products on nutrient cycling have been documented for various species and ecosystems, but temporal variation in these processes is poorly understood. We examined variation in excretion rates of a dominant grazing snail, Elimia clavaeformis, and its contribution to nutrient cycling, over the course of 14 months in a well-studied, low-nutrient stream (Walker Branch, east Tennessee, USA). Biomass-specific excretion rates of ammonium varied over twofold during the study, coinciding with seasonal changes in food availability (measured as gross primary production) and water temperature (multiple linear regression, R ² = 0.57, P = 0.053). The contribution of ammonium excretion to nutrient cycling varied with seasonal changes in both biological (that is, nutrient uptake rate) and physical (that is, stream flow) variables. On average, ammonium excretion accounted for 58% of stream water ammonium concentrations, 26% of whole-stream nitrogen demand, and 66% of autotrophic nitrogen uptake. Phosphorus excretion by Elimia was contrastingly low throughout the year, supplying only 1% of total dissolved phosphorus concentrations. The high average N:P ratio (89:1) of snail excretion likely exacerbated phosphorus limitation in Walker Branch. To fully characterize animal excretion rates and effects on ecosystem processes, multiple measurements through time are necessary, especially in ecosystems that experience strong seasonality.     

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

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

High nutrient availability leads to weaker top‐down control of stream periphyton: Compensatory feeding in Ancylus fluviatilis

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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.     

Mineralisation of dissolved organic matter by heterotrophic stream biofilm communities in a large boreal catchment

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Long‐term responses of leaf litter decomposition to temperature,litter quality and litter mixing in plateau wetlands

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Influence of experimental,environmental, and geographic factors on nutrient‐diffusing substrate experiments in running waters

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Responses of benthic algae to nutrient enrichment in a shallow lake: Linking community production,biomass, and composition

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Biomass loss and change in species dominance shift stream community excretion stoichiometry during severe drought

1. Animals contribute significantly to nutrient cycling through excretion, but most studies consider their effects under relatively benign abiotic conditions. Disturbances such as drought may alter animals’ nutrient contributions through shifts in species composition and biomass. Headwater streams are particularly vulnerable to extreme climate events and thus might show rapid changes in stream biota and their ecosystem effects.
2. We tested how biomass and subsequent ecosystem effects (nutrient cycling) of an intermittent prairie stream community changed during a drought. We quantified the biomass and contributions to nutrient cycling for assemblages comprising fishes, crayfish, and tadpoles in 12 isolated pools over 3 months encompassing the harshest drought on record for Kings Creek, KS, U.S.A. We predicted that macroconsumer biomass would decline with pool surface area and that differences in macroconsumer biomass and taxonomic composition would lead to different contributions of pool assemblages to nutrient cycling.
3. The biomass of pool assemblages declined with decreasing pool size, a pattern apparently driven by mortality, emigration, or metamorphosis. We also observed a change in assemblage structure of drying pools during drought relative to pool size, shifting dominance toward species with more drought-resistant traits. Accordingly, assemblage nitrogen (N) excretion rates declined as pool biomass was reduced, leading to a 58% reduction in N available to epilithic biofilms. Phosphorus (P) excretion rates declined from June to July, but increased in August, as species with high P excretion rates maintained similar proportional biomass and biomass of a non-native fish increased. Molar N:P of pool assemblage excretion declined significantly throughout the drought and coincided with loss of southern redbelly dace (Chrosomus erythrogaster: Cyprinidae).
4. Animal-mediated nutrient cycling was altered by the loss of biomass and stoichiometric traits of taxa that differed in their occurrences and ability to tolerate abiotic conditions during drought. Elevated availability of dissolved N in isolated pools may increase N uptake rates by biofilms during drought conditions, indicating the importance of N excreted by aggregated macroconsumers, especially those with unique stoichiometric traits. While the significance of shifts in the composition of freshwater communities to ecosystems is not entirely known, additional losses in ecosystem function and changes in community structure may follow episodes of severe drought.

     

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.

     

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.     

Effects of stream phosphorus levels on microbial respiration

SUMMARY 1. We examined microbial respiration among streams in lowland Costa Rica comprising a natural phosphorus gradient (5–350 μg SRP L^?1) resulting from variable inputs of solute‐rich (e.g. P, SO₄ and Cl) groundwater. 2. Microbial respiration rates were determined by measuring oxygen change in situ in nine low‐order streams on three substrate types: mixed leaves collected from the stream bottom, conditioned Ficus leaves and sediments. 3. Respiration rates on both leaf types were positively related to phosphorus and negatively related to N : P ratios. Microbial respiration rates on sediments were not related to any of the variables [i.e. soluble reactive phosphorus (SRP), N‐NO₃ and N : P] measured. 4. Respiration rates on newly colonised Ficus leaves formed an asymptotic curve increasing to a plateau, suggesting that saturation with phosphorus occurred at concentrations <15 μg SRP L^?1. 5. To test the hypothesis that phosphorus was the main solute in solute‐rich water that was driving observed differences in microbial respiration rates, we artificially enriched a small stream with phosphorus and measured changes in respiration before and after enrichment. 6. Experimental phosphorus enrichment produced increases in respiration rates similar in magnitude to those observed in the nine streams forming the natural phosphorus gradient, supporting our hypothesis that phosphorus was the major variable driving interstream differences in microbial respiration rates. Respiration rates were higher in this study than those reported for most other tropical streams and rivers with the exception of those reported for tropical Asian streams. 7. Results indicate that variations in phosphorus concentrations can potentially affect patterns of microbial respiration rates at a landscape level via differential inputs of solute‐rich groundwater into streams.     

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.     

Influence of stream flow regime and temperature on growth rate of the riparian tree, Platanus wrightii, in Arizona

1. The growth of riparian trees in semi‐arid regions is influenced by stream flow regime, but the relative importance of base flow and seasonal floods on growth has not been explored. I examined abiotic influences on the growth of Platanus wrightii in four stream reaches in Arizona. All reaches had a bimodal pattern of discharge, but only two had continuous flow throughout the growing season.
2. In two reaches of Sycamore Creek without perennial flow, a large percentage of the annual variation in radial growth rate of P. wrightii was explained by annual and growing season flow rate. Growth was related to these same variables in a perennial reach of Sycamore Creek, but trees maintained higher growth during drought years than they did in the temporary reaches. At Oak Creek, a larger perennial stream, P. wrightii growth showed a bell‐shaped relationship with flow. These data suggest that growth rate is frequently limited by water availability at Sycamore Creek, but not at Oak Creek.
3. At both rivers, much of the annual surface flow occurs as winter floods. Oak Creek, however, maintains a high summer base flow even during years with no floods. Platanus wrightii growth was significantly related to winter flood frequency only at Sycamore Creek. The positive relationship of growth with stream flow and winter flood frequency at Sycamore Creek presumably occurs because the P. wrightii trees are dependent on the winter flows to recharge the shallow alluvial aquifer and to raise the level of ground water within the root zone.
4. Frequent summer floods increased the growth of trees in perennial and non‐perennial reaches alike. At perennial Oak Creek, summer flood frequency was the only variable linearly related to growth of P. wrightii. Summer flood frequency was a significant, but secondary, component of multiple‐regression growth models for trees in the perennial and non‐perennial reaches of Sycamore Creek. Summer floods may stimulate growth, in part, by replenishing limiting nutrients.
5. High temperature was negatively associated with the growth of P. wrightii at Sycamore Creek. The combination of drought and high temperature resulted in very low growth rate.
6. These results have implications for the management of flood and base flow regimes on regulated, diverted and pumped rivers.     

Seasonal changes of C:P ratios of seston, bacteria, phytoplankton and zooplankton in a deep, mesotrophic lake

• 1 The C:P ratios of seston, bacteria, phytoplankton and zooplankton were measured twice a week in situ in mesotrophic, large and deep Lake Constance from April to December 1995. Except for zooplankton, a strong seasonality was exhibited with low C:P ratios during P‐enriched early spring conditions and high values during P‐depleted summer conditions.
• 2 Molar C:P ratios of seston varied between 180:1 and 460:1 demonstrating moderate phosphorus limitation in spring and during the clear‐water phase, and strong limitation for the rest of the season. The sestonic C:P ratio increased significantly during two decades of re‐oligotrophication of Lake Constance, reflecting an enhanced phosphorus limitation of the plankton community in summer. Molar C:P ratios of bacteria and phytoplankton varied seasonally between 50:1 and 130:1 and 180:1 and 500:1, respectively, and indicate carbon or light limitation in winter and phosphorus limitation in summer. Zooplankton had a molar C:P ratio of about 124:115 which was nearly constant throughout the seasons.
• 3 These differences in the C:P ratios of planktonic organisms have direct implications for phosphorus recycling within the food web as C:P ratios of excreta should be highly variable.

     

Nonlinear response of stream ecosystem structure to low‐level phosphorus enrichment

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