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

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

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.     

The influence of ammonium and chloride on potassium and nitrate absorption by barley roots depends on time of exposure and cultivar

Net uptakes of K⁺ and NO₃⁻ were monitored simultaneously and continuously for two barley (Hordeum vulgare) cultivars, Prato and Olli. The cultivars had similar rates of net K⁺ and NO₃⁻ uptake in the absence of NH₄⁺ or Cl⁻. Long-term exposure (over 6 hours) to media which contained equimolar mixtures of NH₄⁺, K⁺, Cl⁻, or NO₃⁻ affected the cultivars very differently: (a) the presence of NH₄⁺ as NH₄Cl stimulated net NO₃⁻ uptake in Prato barley but inhibited net NO₃⁻ uptake in Olli barley; (b) Cl⁻ inhibited net NO₃⁻ uptake in Prato but had little effect in Olli; and (c) NH₄⁺ as (NH₄)₂SO₄ inhibited net K⁺ uptake in Prato but had little effect in Olli. Moreover, the immediate response to the addition of an ion often varied significantly from the long-term response; for example, the addition of Cl⁻ initially inhibited net K⁺ uptake in Olli barley but, after a 4 hour exposure, it was stimulatory. For both cultivars, net NH₄⁺ and Cl⁻ uptake did not change significantly with time after these ions were added to the nutrient medium. These data indicate that, even within one species, there is a high degree of genotypic variation in the control of nutrient absorption.     

AMMONIUM,NITRATE, PHOSPHATE,AND INORGANIC CARBON UPTAKE IN AN OLIGOTROPHIC LAKE: SEASONAL VARIATIONS AMONG LIGHT RESPONSE VARIABLES1

The dependence of substrate saturated uptake of ¹⁵NH₄⁺, ¹⁵NO₃^?, ³²PO₄^3?, and ¹⁴CO₂ on photosynthetic photon flux density (PPFD or photsynthetically active radiation, 400–700 nm) was characterized seasonally in oligotrophic Flathead Lake, Montana. PO₄^3? uptake was not dependent upon PPFD at any time of the year, whereas NH₄⁺, NO₃^?, and CO₂ uptake were consistently dependent on PPFD over all seasons. Maximal rates of NH₄⁺, NO₃^? and CO₂ uptake usually occurred near 40% of surface PPFD, which corresponded to about 5 m in the lake; inhibition was evident at PPFD levels greater than 40%. NH₄⁺, NO₃^? and PO₄^3? were incorporated in the dark at measurable rates most of the year, whereas dark CO₂ uptake was always near 0 relative to light uptake. CO₂ and NO₃^? uptake were more strongly influenced by PPFD than was NH₄^3? uptake. The PPFD dependence of PO₄^3?, NH₄⁺, NO₃^? and CO₂ uptake may affect algal growth and nutrient status by influencing the balance in diel and seasonal C:N:P uptake ratios.     

The effect of land use on dissolved organic carbon and nitrogen uptake in streams

1. Agricultural and urban land use may increase dissolved inorganic nitrogen (DIN) concentrations in streams and saturate biotic nutrient demand, but less is known about their impacts on the cycling of organic nutrients. To assess these impacts we compared the uptake of DIN (as ammonium, NH₄⁺), dissolved organic carbon (DOC, as acetate), and dissolved organic nitrogen (DON, as glycine) in 18 low‐gradient headwater streams in southwest Michigan draining forested, agricultural, or urban land‐use types. Over 3 years, we quantified uptake in two streams in each of the three land‐use types during three seasons (spring, summer and autumn). 2. We found significantly higher NH₄⁺ demand (expressed as uptake velocity, V_f) in urban compared to forested streams and NH₄⁺V_f was greater in spring compared to summer and autumn. Acetate V_f was significantly higher than NH₄⁺ and glycine V_f, but neither acetate nor glycine V_f were influenced by land‐use type or season. 3. We examined the interaction between NH₄⁺ and acetate demand by comparing simultaneous short‐term releases of both solutes to releases of each solute individually. Acetate V_f did not change during the simultaneous release with NH₄⁺, but NH₄⁺V_f was significantly higher with increased acetate. Thus, labile DOC V_f was not limited by the availability of NH₄⁺, but NH₄⁺V_f was limited by the availability of labile DOC. In contrast, neither glycine nor NH₄⁺V_f changed when released simultaneously indicating either that overall N‐uptake was saturated or that glycine and NH₄⁺ uptake were controlled by different factors. 4. Our results suggest that labile DOC and DON uptake can be equivalent to, or even higher than NH₄⁺ uptake, a solute known to be highly bioreactive, but unlike NH₄⁺ uptake, may not differ among land‐use types and seasons. Moreover, downstream export of nitrogen may be exacerbated by limitation of NH₄⁺ uptake by the availability of labile DOC in headwater streams from the agricultural Midwestern United States. Further research is needed to identify the factors that influence cycling of DOC and DON in streams.     

Spatial variability in nutrient concentration and biofilm nutrient limitation in an urban watershed

Nutrient enrichment threatens river ecosystem health in urban watersheds, but the influence of urbanization on spatial variation in nutrient concentrations and nutrient limitation of biofilm activity are infrequently measured simultaneously. In summer 2009, we used synoptic sampling to measure spatial patterns of nitrate (NO₃ ⁻), ammonium (NH₄ ⁺), and soluble reactive phosphorus (SRP) concentration, flux, and instantaneous yield throughout the Bronx River watershed within New York City and adjacent suburbs. We also quantified biofilm response to addition of NO₃ ⁻, phosphate (PO₄ ³⁻), and NO₃ ⁻ + PO₄ ³⁻ on organic and inorganic surfaces in the river mainstem and tributaries. Longitudinal variation in NO₃ ⁻ was low and related to impervious surface cover across sub-watersheds, but spatial variation in NH₄ ⁺ and SRP was higher and unrelated to sub-watershed land-use. Biofilm respiration on organic surfaces was frequently limited by PO₄ ³⁻ or NO₃ ⁻ + PO₄ ³⁻, while primary production on organic and inorganic surfaces was nutrient-limited at just one site. Infrequent NO₃ ⁻ limitation and low spatial variability of NO₃ ⁻ throughout the watershed suggested saturation of biological N demand. For P, both higher biological demand and point-sources contributed to greater spatial variability. Finally, a comparison of our data to synoptic studies of forested, temperate watersheds showed lower spatial variation of N and P in urban watersheds. Reduced spatial variation in nutrients as a result of biological saturation may represent an overlooked effect of urbanization on watershed ecology, and may influence urban stream biota and downstream environments.     

How do chronic nutrient loading and the duration of nutrient pulses affect nutrient uptake in headwater streams?

Our study aimed to analyze the effects of chronic nutrient loading on the capacity of headwater streams to retain phosphorus and ammonium pulses of different duration. For this purpose, we selected nine headwater streams located across a gradient of increasing agricultural land use and eutrophication. In each stream, we performed sequential plateau additions with increasing nutrient concentrations in summer 2015 and instantaneous slug additions in summer 2016 under similar hydrological conditions. We modelled kinetic uptake curves from the slug additions via the Tracer Additions for Spiraling Curve Characterization method and calculated ambient uptake parameters. Ambient uptake rates generally increased (1.4–20.8 µg m^?2 s^?1 for NH₄–N and 0.3–10.3 µg m^?2 s^?1 for SRP, respectively), while ambient uptake velocities decreased from oligotrophic to polytrophic streams (1.8–14.0 mm min^?1 for NH₄–N and 1.6–9.9 mm min^?1 for SRP, respectively). However, correlations between ambient uptake parameters and background concentrations were weak. Concentration-dependent uptake rates followed either a linear or a Michaelis–Menten saturation model, regardless of the degree of nutrient loading. Uptake rate curves showed counter-clockwise hysteresis in oligotrophic streams and clockwise hysteresis in streams of higher trophic states, indicating a reduced significance of hyporheic uptake with increasing nutrient loading. Comparisons of slug and plateau additions revealed that oligotrophic streams were most efficient in uptake during short nutrient pulses, while eutrophic streams profited from longer pulse duration. The results indicate that nutrient uptake is increasingly transport-controlled in polluted streams where increased biofilm thickness and clogging of sediments restrict nutrient transport to reactive sites.     

Inter-annual,Annual, and Seasonal Variation of P and N Retention in a Perennial and an Intermittent Stream

Headwater streams represent the key sites of nutrient retention, but little is known about temporal variation in this important process. We used monthly measurements over 2 years to examine variation in retention of soluble reactive phosphorus (SRP) and ammonium (NH₄⁺) in two Mediterranean headwater streams with contrasting hydrological regimes (that is, perennial versus intermittent). Differences in retention between streams were more evident for NH₄⁺, likely due to strong differences in the potential for nitrogen limitation. In both streams, nutrient-retention efficiency was negatively influenced by abrupt discharge changes, whereas gradual seasonal changes in SRP demand were partially controlled by riparian vegetation dynamics through changes in organic matter and light availability. Nutrient concentrations were below saturation in the two streams; however, SRP demand increased relative to NH₄⁺ demand in the intermittent stream as the potential for phosphorus limitation increased (that is, higher dissolved inorganic nitrogen:SRP ratio). Unexpectedly, variability in nutrient retention was not greater in the intermittent stream, suggesting high resilience of biological communities responsible for nutrient uptake. Within-stream variability of all retention metrics, however, increased with increasing time scale. A review of studies addressing temporal variation of nutrient retention at different time scales supports this finding, indicating increasing variability of nutrient retention with concomitant increases in the variability of environmental factors from the diurnal to the inter-annual scale. Overall, this study emphasizes the significance of local climate conditions in regulating nutrient retention and points to potential effects of changes in land use and climate regimes on the functioning of stream ecosystems.     

Influence of phytoplankton on the response of bacterioplankton growth to nutrient enrichment

1. Laboratory experiments were conducted to test the effect of nutrient enrichment on bacterioplankton growth in the presence and absence of phytoplankton. 2. In one series of experiments, bacterioplankton growth in terms of specific activity [³H-thymidine incorporation (cell number)^?1] was greater in whole lake water samples than in samples from which phytoplankton had been removed by filtration (1.0 μm), regardless of the nutrient enrichments (control, NH⁺₄ plus PO^3-₄ and mannitol). Organic C enhanced bacterioplankton growth in both whole and filtered lake water. 3. In another series of experiments (with the same nutrient enrichments as in the first experiment except that glucose replaced mannitol), bacterioplankton growth in whole lake water enriched with PO^3-₄ plus NH⁺₄ and incubated in the light was greater than in two treatments designed to inhibit photosynthetic activity (+DCMU and dark). Bacterioplankton response to nutrient addition was greatest in the PO^3-₄ plus NH⁺₄ enrichment under all three conditions (light +DCMU, and dark). 4. These results indicate that bacterioplankton growth could be directly limited by inorganic P and N when these elements are in short supply. Enhancement of bacterioplankton growth by phytoplankton occurs only under PO^3-₄ and NH⁺₄ replete environments.     

Nutrient recycling by fish in streams along a gradient of agricultural land use

Nutrient recycling is an essential ecosystem process provided by animals. In many aquatic systems, fish have been identified as important in ecosystem nutrient recycling; however, this importance can vary widely between systems. The factors controlling intersystem variation in animal‐mediated nutrient cycling have rarely been examined and as such it remains unclear what impact human landscape changes will have upon these processes. Here we examined rates of nutrient recycling for temperate stream fish assemblages along a gradient of agricultural land use (proportion cropland in the watershed: 1–59%). We quantified nutrient excretion rates of both ammonium–N (NH₄⁺–N) and phosphate (as soluble reactive phosphate: SRP) for fish assemblages at eight streams in southern Ontario, Canada with species‐specific excretion measurements and quantitative assemblage sampling. For both nutrients, total assemblage excretion exhibited a strong positive relationship with riparian cropland. The distance required for fish assemblages to turn over ambient nutrient pools was shorter for cropland systems, indicating that the relative importance of excreted nutrients was higher in these systems. Based on measured uptake rates of NH₄⁺–N in two streams (one higher cropland and one low cropland) and on modeled uptake rates for all streams, the proportion of ecosystem demand that can be satisfied by excretion is generally higher in the more agricultural streams. These patterns appear to be driven largely by disproportionate increases in fish assemblage biomass with increasing stream nutrient concentrations.     

Amazon deforestation alters small stream structure, nitrogen biogeochemistry and connectivity to larger rivers

Human activities that modify land cover can alter the structure and biogeochemistry of small streams but these effects are poorly known over large regions of the humid tropics where rates of forest clearing are high. We examined how conversion of Amazon lowland tropical forest to cattle pasture influenced the physical and chemical structure, organic matter stocks and N cycling of small streams. We combined a regional ground survey of small streams with an intensive study of nutrient cycling using ¹⁵N additions in three representative streams: a second-order forest stream, a second-order pasture stream and a third-order pasture stream. These three streams were within several km of each other and on similar soils. Replacement of forest with pasture decreased stream habitat complexity by changing streams from run and pool channels with forest leaf detritus (50% cover) to grass-filled (63% cover) channel with runs of slow-moving water. In the survey, pasture streams consistently had lower concentrations of dissolved oxygen and nitrate (NO₃ ^?) compared with similar-sized forest streams. Stable isotope additions revealed that second-order pasture stream had a shorter NH₄ ⁺ uptake length, higher uptake rates into organic matter components and a shorter ¹⁵NH₄ ⁺ residence time than the second-order forest stream or the third-order pasture stream. Nitrification was significant in the forest stream (19% of the added ¹⁵NH₄ ⁺) but not in the second-order pasture (0%) or third-order (6%) pasture stream. The forest stream retained 7% of added ¹⁵N in organic matter compartments and exported 53% (¹⁵NH₄ ⁺?=?34%; ¹⁵NO₃ ^??=?19%). In contrast, the second-order pasture stream retained 75% of added ¹⁵N, predominantly in grasses (69%) and exported only 4% as ¹⁵NH₄ ⁺. The fate of tracer ¹⁵N in the third-order pasture stream more closely resembled that in the forest stream, with 5% of added N retained and 26% exported (¹⁵NH₄ ⁺?=?9%; ¹⁵NO₃ ^??=?6%). These findings indicate that the widespread infilling by grass in small streams in areas deforested for pasture greatly increases the retention of inorganic N in the first- and second-order streams, which make up roughly three-fourths of total stream channel length in Amazon basin watersheds. The importance of this phenomenon and its effect on N transport to larger rivers across the larger areas of the Amazon Basin will depend on better evaluation of both the extent and the scale at which stream infilling by grass occurs, but our analysis suggests the phenomenon is widespread.     

The effect of nutrient concentrations,nutrient ratios and temperature on photosynthesis and nutrient uptake by Ulva prolifera: implications for the explosion in green tides

The green-tide macroalga, Ulva prolifera, was tested in the laboratory to determine its nutrient uptake and photosynthesis under different conditions. In the nutrient concentration experiments U. prolifera showed a saturated uptake for nitrate but an escalating uptake in the tested range for phosphorus. Both N/P and NO₃ ^?/NH₄ ⁺ ratios influenced nutrient uptake significantly (p?<?0.05) while the PSII quantum yield [Y(II)] (p?>?0.05) remained unaffected. The maximum N uptake rate (33.9?±?0.8 μmol g^?1 DW h^?1) and P uptake rate (11.1?±?4.7) was detected at N/P ratios of 7.5 and 2.2, respectively. U. prolifera preferred NH₄ ⁺-N to NO₃ ^?-N when the NO₃ ^?-N/NH₄ ⁺-N ratio was less than 2.2 (p?<?0.05). But between ratios of 2.2 and 12.9, the uptake of NO₃ ^?-N surpassed that of NH₄ ⁺-N. In the temperature experiments, the highest N uptake rate and [Y(II)] were observed at 20 °C, while the lowest rates were detected at 5 °C. P uptake rates were correlated with increasing temperature.     

SHORT TERM UPTAKE OF NUTRIENTS BY ENTEROMORPHA PROLIFERA (CHLOROPHYCEAE)1

Rates of NH₄⁺ and NO₃^? uptake were determined by accumulation of ¹⁵N in plant tissue and by disappearance of nutrient from the medium. Agreement between rates calculated by the two methods was good, averaging 82.7% (SD = 15.8%) and 91.2% (SD = 13.7%) for NH₄⁺ and NO₃^? uptake, respectively. An average of 93.4 and 96.0% of added ¹⁵NH₄⁺ and ¹⁵NO₃^? was recovered from the medium and /or plant tissue at the end of the incubations. Both bacterial uptake and regeneration of NH₄⁺ may contribute to discrepancies between NH₄⁺ uptake rates calculated by ¹⁵N accumulation and disappearance of NH₄⁺ from the medium. The influence of tissue composition on uptake of NH₄⁺, NO₃^? and PO₄^3- by Enteromorpha prolifera (Müller) J. Agardh was examined. For NH₄⁺ uptake, V_max was 188 μmol NH₄^+. g dry wt^?1. h^?1 and K_s ranged from 9.3 to 13.4 μM, but there was no correlation between kinetic parameters and tissue nitrogen content. For NO₃^?, both kinetic parameters were higher for plants with low tissue nitrogen than for plants with high tissue nitrogen. Maximum rates were 169 and 75.4 μmol NO₃^?. g dry wt^?1. h^?1, and K_s was 13.3 and 2.31 μM for low and high tissue nitrogen plants, respectively. Estimates of uptake in the field suggested that NH₄⁺ accounted for 65% and NO₃^? for up to 35% of total nitrogen uptake during the summer. Nutrient uptake rates of field-collected plants also indicated that E. prolifera in Yaquina Bay, Oregon was not likely to have been nitrogen-limited, but may have been phosphorus-limited.     

Simulation study of nutrient uptake by plants from soilless cultures as affected by salinity buildup and transpiration

Soilless plant growth systems are widely used as a means to save irrigation water and to reduce groundwater contamination. While nutrient concentrations in the growth medium are depleted due to uptake by the plants, salinity and toxic substances accumulate due to transpiration. A theoretical model is suggested, to simulate nutrient uptake by plants grown in soilless cultures with recycled solutions. The model accounts for salinity accumulation with time and plant growth, and its effects on uptake of the different nutrients by means of interaction with Na and Cl ions. The sink term occurs due to uptake by a growing root system. Influx as a function of the ion concentration is according to Michaelis–Menten active mechanisms for K⁺, NO₃ ^–-N, NH₄ ⁺-N, PO₄-P, Ca²⁺, Mg²⁺ and SO₄ ^2-, whose influx parameters are affected by Na and Cl^–, but not with time (age). Sodium influx is passive above a critical concentration. Sum of cations–anions concentrations is balanced by Cl^– to maintain electro-neutrality of the growth solution. Salinity (by means of Na concentration) suppresses root and leaf growth, which further effect uptake and transpiration. The model accounts for instantaneous transpiration losses, during daytime only and its effect on uptake of nutrients and plant development due to salt accumulation. The model was tested against NO₃ ^– and K⁺ uptake by plants associated with cumulative transpiration and with different NaCl salinity levels. Deviations from observed K⁺ uptake should be attributed to the salinity tolerance of the plants. In a study with data obtained from published literature, the model indicated that nutrient depletion and salinity buildup might be completely different with fully grown-up plants (that do not grow) and plants that grow with time. Depletion of different nutrients are according to their initial concentration and plant uptake rate, but also affected by their interactions with Na and Cl ions.     

Forest Age Influences In-stream Ecosystem Processes in Northeastern US

A disturbance or natural event in forested streams that alter available light can have potential consequences for nutrient dynamics and primary producers in streams. In this study, we address how functional processes (primary production and nutrient uptake) in stream ecosystems respond to changes in forest canopy structure. We focus on differences in incoming irradiance, nutrient uptake (NO₃, NH₄, and PO₄) and open-channel metabolism seasonally in 13 forested streams that drain forests with different canopy structures (10 to >300 years old) in the northeastern United States. Light irradiance was related to forest age in a U-shaped pattern, with light being the greatest in both young open forests (<50 years old) and older growth forests (>245 years old), whereas the darkest conditions were found in the secondary growth middle-aged forests (80–158 years old). Streams that had adjacent open or old-growth riparian forest had similar conditions with greater standing stock biofilm biomass (chl a), and elevated ER in October compared to streams with middle-aged riparian forests. Compared to all sites, streams with old-growth riparian forest had the greatest in-stream primary production rates (GPP) and elevated background nutrient concentrations, and to a lesser degree, increased nutrient retention and uptake (V _f). Streams draining older forests tended to be more productive and retentive than middle-aged forests, likely due to increased light availability and the age and structure of surrounding forest canopies. Middle-aged forests had the least variation in response variables compared to streams in young and old-growth riparian forests, likely a result of uniform canopy conditions. As the structure of widespread middle-aged forests in NE US is altered by loss of specific tree species, climate change, and/or human activity, it will impact in-stream production and nutrient dynamics and may ultimately alter nutrient loading in downstream catchments.     

Ecological indicators of nutrient enrichment, freshwater wetlands, Midwestern United States (U.S.)

Vegetation and soil indicators of nutrient condition were evaluated in 30 wetlands, 10 each in 3 Nutrient Ecoregions (NE) (VI-Corn Belt and Northern Great Plains, VII-Mostly Glaciated Dairy Region, IX-Temperate Forested Plains and Hills) of the Midwestern United States (U.S.) to identify robust indicators for assessment of wetland nutrient enrichment and eutrophication. Nutrient condition was characterized by surface water inorganic N (NH₄-N, NO₃-N) and P (PO₄-P) concentrations measured seasonally for 1 year, plant available and total soil N and P, and aboveground biomass, leaf N and P and species composition of emergent vegetation measured at the end of the growing season. Aboveground biomass, nutrient uptake and species composition were positively related to surface water NH₄-N (N) but not to PO₄-P or NO₃-N. Aboveground biomass and biomass of aggressive species, Typha spp. plus Phalaris arundinacea, increased asymptotically with surface water N whereas leaf P, senesced leaf N and senesced leaf P increased linearly with N. And, species richness declined with surface water N. Soil total P was positively related to surface water PO₄-P but it was the only soil indicator related to wetland nutrient condition. Individual regressions for each NE generally were superior to a single regression for all NEs. In NE VI (Corn Belt), few indicators were related to surface water N because of the high degree of anthropogenic disturbance (85% of the landscape is cleared) as compared to NEs VII and IX (24–53% cleared). Of the indicators evaluated, stem height (r² = 0.42 for all NEs, r² = 0.56 for NE VII + IX) and percent biomass of aggressive species, Typha spp. plus Phalaris, (r² = 0.46 for all NEs, r² = 0.54 for NE VII + IX), were the best predictors of wetland nutrient enrichment. Vegetation-based indicators are a promising tool for assessment of wetland nutrient condition but they may not be effective in NEs where landscape disturbance is intense and widespread.     

Effects of pH,phosphate and ammonia on the rate of uptake of nitrate and ammonia by freshwater phytoplankton

The effect of phosphate (PO₄ ⁺³) and pH in regulating nitrate (NO₃) and ammonia (NH₃ ⁺) uptake by phytoplankton was investigated in two Oklahoma lakes using ¹⁵N tracers. Addition of PO₄ ⁺³ above ambient concentrations had a negligible effect on the rate of uptake of NO₃ ^– or NH₃ ⁺. Manipulation of pH of lake water had little effect on uptake of either NO₃ ^– or NH₃ ⁺. A correlation analysis suggested that NO₃ ^– is not used by phytoplankton when NH₃ ⁺ concentrations exceed about 210 µg NH₃ ⁺-N(1)^–1.