Effects of moderate ammonium enrichment on three submersed macrophytes under contrasting light availability

1. Increased ammonium concentrations and decreased light availability in a water column have been reported to adversely affect submersed vegetation in eutrophic waters worldwide. 2. We studied the chronic effects of moderate enrichment (NH₄–N: 0.16–0.25 mg L^?1) on the growth and carbon and nitrogen metabolism of three macrophytes (Ceratophyllum demersum, Myriophyllum spicatum and Vallisneria natans) under contrasting light availability in a 2‐month experiment. 3. The enrichment greatly increased the contents of free amino acids and nitrogen in the shoot / leaf of the macrophytes. This indicates that was the dominant N source for the macrophytes. 4. Soluble carbohydrate contents remained relatively stable in the shoot / leaf of the macrophytes irrespective of the treatments. Under ambient light, the starch contents in the shoot / leaf of C. demersum and M. spicatum increased with enrichment, whereas V. natans did not exhibit any change. The starch contents decreased in C. demersum, increased in M. spicatum and remained unchanged in V. natans after the combined treatment of enrichment and reduced light. 5. The enrichment did not affect the growth of the three macrophytes under the ambient light. However, it did suppress the growth of C. demersum and M. spicatum under the reduced light. The results indicate that a moderate enrichment was not directly toxic to the macrophytes although it might change their viability in eutrophic lakes in terms of the carbon and nitrogen metabolism.     

Planktonic phosphorus pool sizes and cycling efficiency in coastal and interior British Columbia lakes

1. Limnologists have long acknowledged the importance of phosphorus (P) in determining the organism biomass and productivity of lake ecosystems. Despite a relatively large number of studies that have examined P cycling in lake ecosystems, there remain several substantial methodological issues that have impeded our understanding of P cycling in limnetic plankton communities. Two critical issues confronting ecologists are (1) a lack of precise measurements of the dissolved inorganic phosphorus (PO) and (2) accurate or complete measurements of dissolved P regeneration rates by plankton communities. 2. Here, we examine patterns of epilimnetic planktonic P pool sizes and turnover rates in eight lakes in British Columbia, Canada over a 2‐year period. We determine the concentrations and turnover times of P in various planktonic compartments (dissolved and various planktonic size fractions), using recently developed methods for estimating phosphate concentration and planktonic regeneration rates. 3. The pico‐ and nanoplankton size fraction (0.2–20 μm) played a central role in planktonic P cycling in lakes examined by this study. On average across lakes, pico‐ and nanoplankton contained >60% of the planktonic P, accounted for >90% PO uptake, and contributed 50% of the plankton community dissolved P regeneration rate. 4. PO concentrations determined by steady state bioassays (ssPO) were extremely low (87–611 pmol L⁻¹) and were 2–3 orders of magnitude less than simultaneously measured colorimetric soluble reactive phosphorus estimates. Lake ssPO concentrations increased linearly with total phosphorus (TP), and the slope of this relationship was approximately 1, indicating that PO remained a consistent proportion of the TP pool across a range of TP concentrations. 5. Turnover rates of the total planktonic P pool and the <20 μm pool became more rapid with increasing lake TP, indicating that, according to this metric, planktonic P cycling efficiency increased with TP concentrations. We also detected a significant relationship between particulate phosphorus (PP) <20 μm turnover time and seston N : P ratios, with PP <20 μm turnover times becoming slower with increasing seston N : P. These findings suggest that long‐standing conceptual models of nutrient cycling that predict slower cycling rates and decreasing cycling efficiency with increasing TP concentrations require further empirical examination. We postulate that patterns in lake P turnover and cycling efficiency are a result of complex interactions between plankton biomass and composition, and the ratios of multiple nutrients (C, N, P), rather than solely a function of the TP pool.     

Relationships between microcystins and environmental parameters in 30 subtropical shallow lakes along the Yangtze River, China

1. A survey of 30 subtropical shallow lakes in the middle and lower reaches of the Yangtze River area in China was conducted during July–September in 2003–2004 to study how environmental and biological variables were associated with the concentration of the cyanobacterial toxin microcystin (MC). 2. Mean MC concentration in seasonally river‐connected lakes (SL) was nearly 33 times that in permanently river‐connected lakes (RL), and more than six times that in city lakes (NC) and non‐urban lakes (NE) which were not connected to the Yangtze River. The highest MC (8.574 μg L^?1) was detected in Dianshan Lake. 3. MC‐RR and MC‐LR were the primary toxin variants in our data. MC‐RR, MC‐YR and MC‐LR were significantly correlated with Chl a, biomass of cyanobacteria, Microcystis and Anabaena, indicating that microcystins were mainly produced by Microcystis and Anabaena sp. in these lakes. 4. Nonlinear interval maxima regression indicated that the relationships of Secchi depth, total nitrogen (TN) : total phosphorus (TP) and NH with MC were characterised by negative exponential curves. The relationships between MC and TN, TP, NO + NO were fitted well with a unimodal curve. 5. Multivariate analyses by principal component and classifying analysis indicated that MC was mainly affected by Microcystis among the biological factors, and was closely related with temperature among physicochemical factors.     

Distribution and regulation of urea in lakes of central North America

1. Urea accounts for ～50% of global nitrogen (N)‐based fertiliser; however, little is known of the factors regulating its distribution and abundance in freshwaters. Improved understanding of urea biogeochemistry is essential because its use as fertiliser is expected to double by 2050 and because pollution with urea can promote outbreaks of toxic cyanobacteria in phosphorus (P)‐rich lakes in regions with intensive agricultural or urban development. 2. Biweekly measurements of urea concentration and diverse limnological variables (water chemistry, hydrology, algae, zooplankton) were taken during two summers (2008, 2009) in a chain of seven productive lakes within a 52 000‐km² catchment in central Canada to quantify environmental and anthropogenic correlates of temporal and spatial patterns of urea occurrence. 3. Mean (±SD) urea concentrations varied between 29 ± 14 and 132 ± 65 μg N L^?1, generally increased from headwater to downstream sites and represented 10–50% of bioavailable N (as sum of , and urea). Principal components analysis demonstrated that urea concentrations were elevated in agriculturally impacted lakes with abundant dissolved organic and inorganic nutrients (N, P, C) and low O₂ concentrations, but were not correlated consistently with plankton abundance or community composition. Urea concentrations were more than twofold greater in lakes receiving N from cities than in agriculturally affected basins, despite low summer concentrations of urea in tertiary‐treated urban effluent (c. 50% of lake values). Multiple regression models evaluated using Akaike Information Criterion showed that mean water‐column O₂ concentration was the single best predictor of in situ urea concentrations (r² = 0.91, P = 0.002), but that urea concentrations were also correlated significantly with changes in longitudinal position and Secchi depth and with concentrations of , non‐urea dissolved organic N (DON) and dissolved inorganic carbon. 4. Additional seasonal surveys of up to 69 closed‐basin lakes within a 100 000‐km² region during 2004 and 2008 revealed that urea was abundant in 100% of measured sites and exhibited concentrations (81 ± 48 μg N L^?1) similar to those observed in lakes with surface drainage (58 ± 38 μg N L^?1). Further, non‐urea DON accounted for 50–99% of the total dissolved N pool in both open‐ and closed‐basin lakes. 5. When combined with an extensive literature review and previous mass‐budget analyses of the study lakes, these findings allowed the development of a first‐generation model of the mechanisms regulating urea content of P‐rich lakes of central North America. In this model, water‐column concentrations of urea are predicted to be regulated mainly by algal decomposition in anoxic environments (sediments, hypolimnion), followed by redistribution into surface waters. Consequently, anthropogenic activities can increase the urea content of lakes by stimulating primary production, sedimentation and deepwater anoxia and by increasing influx of undegraded urea from agricultural and urban sources.     

Phosphorus limitation of the freshwater benthic diatom Didymosphenia geminata determined by the frequency of dividing cells

1. Unlike other nuisance algal species, the freshwater benthic diatom Didymosphenia geminata typically forms blooms in low‐nutrient rivers. The negative association between D. geminata blooming behaviour and nutrient levels appears at both catchment and smaller scales. We conducted a series of trials in streamside experimental channels colonised with D. geminata using water from the D. geminata‐affected, oligotrophic Waitaki River, South Island, New Zealand to determine how elevated nitrate and phosphate concentrations affected D. geminata cell division. Because D. geminata blooms are typically most pronounced in unshaded waters, we also investigated the growth response to shading. In all experiments, we used the frequency of dividing cells (FDC) as a metric of cell division. 2. Concentrations of nitrate and dissolved reactive phosphorus (DRP) in the Waitaki River were very low (4 mg m^?3‐N and <1 mg m^?3 DRP). In pilot trials, substrata colonised by D. geminata were subjected to enrichment by either switching the water source toN‐ and P‐rich spring water or by adding a stock solution. Both trials resulted in periods of rapid cell division lasting at least 8 days. 3. Experimental addition of alone triggered an initial cell division which was not sustained. However, addition of alone or together with resulted in prolonged elevation in cell division indicating that the cell division rate was P‐limited. 4. Reduced light levels resulted in decreased FDC in D. geminata in both ambient and N, N + P and P‐enriched river water. 5. Stimulation of D. geminata division rate by addition of above ambient levels confirms that, while blooming behaviour is often associated with oligotrophic rivers, the cells divide faster with greater levels of phosphorus enrichment.     

Microbial activities in the burrow environment of the potamal mayfly Ephoron virgo

1. The impact of burrowing larvae of Ephoron virgo (Ephemeroptera, Polymitarcidae) on sediment microbiology has not been previously investigated because of difficulties in sampling the sediment of large rivers under in situ conditions. Therefore, we conducted experiments in the on‐ship Ecological Rhine Station of the University of Cologne (Germany), in which ambient conditions of the River Rhine can be closely mimicked. 2. In two consecutive seasons, experimental flow channels were stocked with Ephoron larvae and continuously supplied with water taken directly from the River Rhine. Sediment from the immediate vicinity of Ephoron burrows (i.e. U‐shaped cavities reaching 10–80 mm deep into the sediment) and bulk sediment samples were analysed for (i) particulate organic matter content, (ii) microscale in situ distribution of O₂, NO, and NH, and (iii) potential activities of exoenzymes. 3. Sediment surrounding the Ephoron burrows had markedly higher organic matter contents and exoenzyme activities compared with the bulk sediment. Microsensor measurements demonstrated that local O₂ and NO penetration into the sediment were greatly enhanced by larval ventilation behaviour. Volumetric O₂ and NO turnover rates that were calculated from steady state concentration profiles measured directly in the burrow lining were considerably higher than at the sediment surface. 4. In the sediment of the fast flowing River Rhine Ephoron burrows are preferential sites of organic matter accumulation and dissolved oxidant penetration. Our data suggest that the burrows are surrounded by a highly active microbial community that responds to the inputs from the water column with elevated O₂ and NO turnover, and release of exoenzymes into the sediment pore water. Especially during periods of mass occurrence, the larvae of E. virgo may thus significantly contribute (i) to the ecological connection between the water column and the sediment and (ii) to biogeochemical processing of organic matter in the riverbed.     

Phosphorus dynamics in Danish lakes and the implications for diatom ecology and palaeoecology

SUMMARY 1. Seasonal phosphorus variability may be considerable in eutrophic lakes and patterns are influenced by internal, as well as external, nutrient loading. The strong seasonality of planktonic diatoms, with main growth periods in the spring and also in the autumn in meso‐ to eutrophic lakes, means that the measure of total phosphorus (TP) that is most relevant for diatom ecology may not be adequately assessed by single or few measurements during the year. 2. The diatom species assemblage of surface sediments can be used to infer in‐lake nutrient concentrations. Weighted averaging (WA) and weighted averaging partial least squares (WAPLS) regression and calibration models for diatoms and annual mean TP were developed for a 29‐site data set of Danish lakes based on (a) all diatom species and (b) planktonic species only. Jack‐knifed error statistics were: =0.37, root mean squared error of prediction (RMSEP)=0.28 log₁₀μg TP L^?1 and mean bias=0.04 log₁₀μg TP L^?1 for the WAPLS 2‐component model based on all species; =0.23, RMSEP= 0.32log₁₀μg TP L^?1 and mean bias=0.07 log₁₀μg TP L^?1 for the WA model with tolerance downweighting based on planktonic species only. These are comparable with similar, published data sets. 3. A subset of 23 sites was used to develop models based on seasonal TP measurements. Mean spring TP concentrations gave only slightly improved RMSEP values for models based on all species and plankton‐only (0.24 log₁₀μg TP L^?1 and 0.29 log₁₀μg TP L^?1, respectively). 4. Weighted averaging models derive environmental optima, for individual species, which are not necessarily of ecological relevance. However, good water chemistry data are required to model species' responses adequately and to develop calibration data sets.     

Effects of irradiance on diel and seasonal patterns of nutrient uptake by stream periphyton

1. Irradiance strongly affects the abundance of stream periphyton communities that in turn influence patterns of instream nutrient uptake. We examined relationships between irradiance and periphyton nutrient uptake taking into account diel and seasonal variation in ambient irradiance. 2. Uptake of dissolved N, P and C by periphyton as areal uptake (U) and demand (V_f) was determined under 11 irradiance levels (0–100% of ambient conditions) using shallow stream‐side experimental channels. Experiments were conducted once per season over one annual cycle with both day and night uptake rates assessed, together with periphyton biomass and autotrophic production rates. 3. No consistent diel variation in areal uptake or demand was detected for the predominant inorganic or total dissolved nutrients even at the highest irradiances. Lack of variation may indicate nutrient limitation, with photosynthetic sequestration and storage of C during the day for subsequent utilisation at night. Alternatively, oxygen consumption by photoautotrophs at night may stimulate compensatory heterotrophic uptake (e.g. denitrification). 4. In all seasons, release of dissolved organic N was detected during the day but to a lesser extent at night. This was not directly related to irradiance levels, indicating that heterotrophic metabolism (e.g. microbial decomposition) contributes to this phenomenon. 5. Areal uptake and demand for the predominant inorganic and total dissolved nutrients increased in response to increasing irradiance in some or all seasons, but rates were typically higher during the spring and summer. Saturation of areal uptake and demand at elevated irradiances was evident during the spring. demand was also saturated at higher irradiances in the summer and autumn. Maximum demand was comparable during spring and summer, but saturation occurred at lower irradiance in summer (24 h average 135–145 μmol m^?2 s^?1) relative to spring (312–424 μmol m^?2 s^?1), indicating more efficient nutrient uptake in summer. Higher total periphyton biomass in summer, but comparable autotrophic biomass (chlorophyll a), implies that heterotrophic metabolism may contribute to this greater efficiency. In spring, autotrophic biomass peaked at an irradiance level of 225 μmol m^?2 s^?1, also suggesting a role for heterotrophic metabolism in demand at higher irradiances. 6. The results of this study show that irradiance levels exert a strong influence on the nature and quantity of instream nutrient uptake with N demand saturated at elevated irradiance levels during the spring, summer and autumn. Our results also suggest that heterotrophic metabolism makes a measurable contribution to instream nutrient uptake even under higher irradiances that favour autotrophic activity.     

Distribution of benthic diatoms in U.S. rivers in relation to conductivity and ionic composition

• 1 We quantified the relationships between diatom relative abundance and water conductivity and ionic composition, using a dataset of 3239 benthic diatom samples collected from 1109 river sites throughout the U.S.A. [U.S. Geological Survey National Water‐Quality Assessment (NAWQA) Program dataset]. This dataset provided a unique opportunity to explore the autecology of freshwater diatoms over a broad range of environmental conditions.
• 2 Conductivity ranged from 10 to 14 500 μS cm^?1, but most of the rivers had moderate conductivity (interquartile range 180–618 μS cm^?1). Calcium and bicarbonate were the dominant ions. Ionic composition, however, varied greatly because of the influence of natural and anthropogenic factors.
• 3 Canonical correspondence analysis (CCA) and Monte Carlo permutation tests showed that conductivity and abundances of major ions (HCO + CO, Cl^?, SO, Ca²⁺, Mg²⁺, Na⁺, K⁺) all explained a statistically significant amount of the variation in assemblage composition of benthic diatoms. Concentrations of HCO + CO and Ca²⁺ were the most significant sources of environmental variance.
• 4 The CCA showed that the gradient of ionic composition explaining most variation in diatom assemblage structure ranged from waters dominated by Ca²⁺ and HCO + CO to waters with higher proportions of Na⁺, K⁺, and Cl^?. The CCA also revealed that the distributions of some diatoms correlated strongly with proportions of individual cations and anions, and with the ratio of monovalent to divalent cations.
• 5 We present species indicator values (optima) for conductivity, major ions and proportions of those ions. We also identify diatom taxa characteristic of specific major‐ion chemistries. These species optima may be useful in future interpretations of diatom ecology and as indicator values in water‐quality assessment.

     

The dual influences of dissolved organic carbon on hypolimnetic metabolism: organic substrate and photosynthetic reduction

We investigated the effect of dissolved organic carbon (DOC) on hypolimnetic metabolism (accumulation of dissolved inorganic carbon (DIC) and methane (CH₄)) in 21 lakes across a gradient of DOC concentrations (308 to 1540 mol C L^–1). The highly colored nature of the DOC in these lakes suggests it is mostly of terrestrial origin. Hypolimnetic methane accumulation was positively correlated with epilimnetic DOC concentration (Spearman rank correlation = 0.67; p < 0.01), an indicator of allochthonous DOC inputs, but not with photic zone chlorophyll a concentration (Spearman rank correlation = 0.30; p = 0.22). Hypolimnetic DOC concentrations declined in 19 of 21 lakes during the stratified period at rates that ranged from 0.06 to 53.9 mmol m^–2 d^–1. The hypolimnetic accumulation of DIC + CH₄ was positively correlated with, and, in most cases of comparable magnitude to, this DOC decline suggesting that DOC was an important substrate for hypolimnetic metabolism. The percentage of surface irradiance reaching the thermocline was lower in high DOC lakes (0.3%) than in low DOC lakes (6%), reducing hypolimnetic photosynthesis (as measured by the depth and magnitude of the deep dissolved oxygen maxima) in the high DOC lakes. In June, the hypolimnia of lakes with < 400 mol L^–1 DOC had high concentrations of dissolved oxygen and no CH₄, while the hypolimnia of lakes with DOC > 800 mol L^–1 were completely anoxic and often had high CH₄ concentrations. Thus, DOC affects hypolimnetic metabolism via multiple pathways: DOC was significant in supporting hypolimnetic metabolism; and at high concentrations depressed photosynthesis (and therefore oxygen production and DIC consumption) in the hypolimnion.     

Nitrogen cycling and dynamics in a macrophyte‐rich stream as determined by a release

1. A tracer release study was conducted in a macrophyte‐rich stream, the River Lilleaa in Denmark. The objectives of the study were to compare uptake rates per unit area of by primary producers and consumers in macrophyte and non‐macrophyte habitats, estimate whole‐stream uptake rates of and compare this to other stream types, and identify the pathways and estimate the rate at which enters the food web in macrophyte and non‐macrophyte habitats. 2. Macrophyte habitats had four times higher primary uptake rates and an equal uptake rate by primary consumers per unit habitat area as compared to non‐macrophyte habitats. These rates represent the lower limit of potential macrophyte effects because the rates will be highly dependent on macrophyte bed height and mean bed height in the River Lilleaa was low compared to typical bed heights in many lowland streams. Epiphytes accounted for 30% of primary uptake in macrophyte habitats, illustrating a strong indirect effect of macrophytes as habitat for epiphytes. N flux per unit habitat area from primary uptake compartments to primary consumers was four times lower in macrophyte habitats compared to non‐macrophyte habitats, reflecting much greater biomass accrual in macrophyte habitats. Thus, we did not find higher N flux from macrophyte habitats to primary consumers compared to non‐macrophyte habitats. 3. Whole‐stream uptake rate was 447 mgN m^?2 day^?1. On a habitat‐weighted basis, fine benthic organic matter (FBOM) accounted for 72% of the whole‐stream uptake rate, and macrophytes and epiphytes accounted for 19 and 8%, respectively. 4. We had expected a priori relatively high whole‐stream N uptake in our study stream compared to other stream types mainly due to generally high biomass and the macrophyte’s role as habitat for autotrophic and heterotrophic organisms, but our results did not confirm this. In comparison with other release study streams, we conclude that nutrient concentration is the overall controlling factor for N uptake rates across streams, mostly as a result of high biomass of primary uptake compartments in streams with high nutrient concentrations in general and not in macrophyte streams in particular. 5. Our results indicate that macrophytes play an important role in the longer‐term retention of N and thus a decrease in net downstream transport during the growing season compared to streams without macrophytes, through direct and indirect effects on the stream reach. Direct effects are high uptake efficiency, low turnover rate (partly due to no direct feeding on macrophytes) and high longevity. An indirect effect is increased sedimentation of FBOM in macrophytes compared to non‐macrophyte habitats and streams which possibly also increase denitrification. Increased retention with macrophyte presence would decrease downstream transport during the growing season and thus the N loading on downstream ecosystems.     

Influence of internal loading on phosphorus concentration in shallow lakes before and after reduction of the external loading

An analysis of data from 49 shallow lakes showed, that the parameters of empirical models between phosphorus loading and concentration in the lake (e.g. Vollenweider type of relations) differ significantly for lakes without or with a reduced external loading. For lakes without a reduction of the external loading the summer phosphorus concentration is determined by the external phosphorus loading and the hydraulic loading. For these lakes the classical models suffice; deviations between calculations and measurements are partly due to errors made in the determination of the loading.In contrast, for lakes where the external loading was reduced, the measured internal loading explains most of the variation in the summer lake concentration. The external loading is of minor importance and the classical models cannot be applied. The internal loading measured before reduction of the external loading is not useful in predicting the concentration afterwards. Instead of the internal loading, the sediment composition can be used. The advantage of using sediment composition is that these variables are easier to determine and vary less in time. The most promising variable is the ratio between total P and total Fe in the sediment.Abbreviations: Q_s hydraulic loading (m y^-1) - hydraulic retention time (y) - L_ext external phosphorus loading (gP m^-2 y^-1) - L_int internal phosphorus loading (gP m^-2 y^-1) - P_lake phosphorus concentration in the lake (gP m^-3) - P_inlet phosphorus concentration in the inlet water (gP m^-3) - P_sed phosphorus content on the sediment (gP kg^-1 d.w.) - Fe_sed iron content of the sediment (gFe kg^-1 d.w.) - Y dependent variable multiple regression calculations - X1, X2 independent variables multiple regressions calculations - a, a1 constants - a2, b constants     

Release rates and potential fates of nitrogen and phosphorus from sediments in a eutrophic reservoir

1. Nutrients released from lake sediments can influence water column nutrient concentrations and planktonic productivity. We examined sediment nutrient release [soluble reactive phosphorus (SRP) and ammonia (NH)] at two sites in a eutrophic reservoir (Acton Lake, OH, U.S.A.) that differed in physical mixing conditions (a thermally stratified and an unstratified site). 2. Sediment nutrient release rates were estimated with three methods: sediment core incubations, seasonal in situ hypolimnetic accumulation and a published regression model that predicted sediment phosphorous (P) release rate from sediment P concentration. All three methods were applied to the deeper stratified site in the reservoir; however, we used only sediment core incubations to estimate SRP and NH release rates at the shallow unstratified site because of the lack of thermal stratification. We also compared the total P concentration (TP_S) of sediments and the concentration of P in various sediment fractions at both sites. 3. Anoxic sediments at the stratified site released SRP at rates more than an order of magnitude greater than oxic sediments at the shallow unstratified site. However, P accumulated in the hypolimnion at much lower rates than predicted by sediment core incubations. In contrast, NH was released at similar rates at both sites and accumulated in the hypolimnion at close to the expected rate, indicating that P was ‘lost’ from the hypolimnion through biogeochemical pathways for P, such as precipitation with inorganic material or biological uptake and sedimentation. 4. TP_S was significantly greater at the deeper stratified site and organically bound P accounted for >50% of TP_S at both sites. 5. We examined the magnitude of SRP fluxes into the study reservoir in 1996 by comparing the mean summer daily SRP fluxes from anaerobic sediments, aerobic sediments, stream inflows and gizzard shad excretion. While the SRP release from anaerobic sediments was high, we hypothesise that little of this SRP gained access to the epilimnion in mid‐summer. SRP flux to the reservoir from aerobic sediments was less than from gizzard shad excretion and streams. Large interannual variability in thermocline stability, gizzard shad biomass and stream discharge volumes, will affect SRP loading rates from different sources in different years. Therefore, construction of P budgets for different years should account for interannual variation in these parameters.     

Are geothermal streams important sites of nutrient uptake in an agricultural and urbanising landscape (Rotorua,New Zealand)?

1. Lakes in the Rotorua region of New Zealand are affected by eutrophication from urbanisation and agricultural land use. Some lake tributaries contain geothermally influenced waters, and it is currently unknown whether geothermal tributaries are active sites of nutrient cycling or represent point sources of nutrients to the lakes. 2. Using government data sets, we characterised the physicochemical conditions of geothermal and non‐geothermal streams. We then measured ecosystem metabolism and reach‐scale uptake of nitrate (), ammonium () and phosphate () in summer 2010 (n = 8 streams). Finally, we used government data to compare annual nutrient flux from geothermal and non‐geothermal surface water inputs to Lake Rotoiti. 3. As expected, geothermal streams had higher temperature, conductivity and nutrient concentrations and lower pH. However, primary production, community respiration and uptake rates in geothermal streams were not different from those in their non‐geothermal counterparts. Uptake rates of were higher in geothermal streams, and uptake was below detection in geothermal streams, probably due to the saturation by naturally high concentrations. 4. A comparison of Lake Rotoiti inputs suggested that geothermal streams are not significant sources of and , while geothermal inputs of represent an average of 46% of total flux from Lake Rotoiti tributaries. 5. Despite their high temperature and low pH, geothermal streams are active sites of photosynthesis, respiration and and cycling, indicating dynamic biofilm communities. 6. Management options for geothermal streams, if any, should focus on retention (e.g. uptake or coupled nitrification and denitrification) but could prove challenging given the persistent, naturally occurring high flux.     

Elevated methylmercury in High Arctic Daphnia and the role of productivity in controlling their distribution

Mercury is a contaminant of concern in polar regions due to long‐range atmospheric transport of this metal from southern latitudes followed by intense deposition on snow. We surveyed zooplankton in 16 lakes and ponds in the Canadian Arctic Archipelago (74–76°N) to determine methylmercury (MeHg) content and the role of environmental characteristics and taxonomic composition on accumulation processes. Zooplankton communities containing Daphnia (mainly D. middendorffiana) had on average five times the MeHg content of copepod‐dominated communities. The percent biomass of Daphnia best explained MeHg variation in bulk zooplankton compared with water chemistry and morphometric variables. Water‐column concentrations of MeHg were low at most study sites (mainly ≤0.07 ng L⁻¹), and Daphnia strongly bioaccumulated mercury through species‐specific processes. As we observed Daphnia in more productive water bodies (i.e., ponds, a eutrophied lake), we then tested the role of productivity in determining the distribution of this keystone herbivore using a broad‐scale literature dataset of 47 High Arctic lakes (65–77°N). Daphnia density was positively related to the amount of organic carbon in the water column in both dissolved and particulate fractions [dissolved organic carbon (DOC) partial , P < 0.001; particulate organic carbon (POC) partial , P=0.032]. The strong influence of DOC suggests that bacterial production is an important energy source for Arctic Daphnia. Our findings indicate that productivity influences the MeHg content of zooplankton communities through its control of species composition; specifically, low productivity limits the presence of mercury‐rich Daphnia in many copepod‐dominated lakes of the High Arctic. Aquatic productivity is expected to increase with climate warming, and we present a conceptual model that predicts how environmental drivers could extend the distribution of Daphnia in lakes and alter the movement of mercury in food webs of the Canadian High Arctic.     

Regulation of nitrous oxide emission associated with benthic invertebrates

1. A number of freshwater invertebrate species emit N₂O, a greenhouse gas that is produced in their gut by denitrifying bacteria (direct N₂O emission). Additionally, benthic invertebrate species may contribute to N₂O emission from sediments by stimulating denitrification because of their bioirrigation behaviour (indirect N₂O emission). 2. Two benthic invertebrate species were studied to determine (i) the dependence of direct N₂O emission on the preferred diet of the animals, (ii) the regulation of direct N₂O emission by seasonally changing factors, such as body size, temperature and availability and (iii) the quantitative relationship between direct and indirect N₂O emission. 3. Larvae of the mayfly Ephemera danica, which prefer a bacteria‐rich detritus diet, emitted N₂O at rates of up to 90 pmol Ind.^?1 h^?1 under in situ conditions and 550 pmol Ind.^?1 h^?1 under laboratory conditions. In contrast, larvae of the alderfly Sialis lutaria, which prefer a bacteria‐poor carnivorous diet, emitted N₂O at invariably low rates of 0–20 pmol Ind.^?1 h^?1. The N₂O emission rate of E. danica larvae was positively correlated with seasonally changing factors (body size, temperature and availability). Direct N₂O emission by E. danica larvae was limited by low temperature in winter, larval development in spring and low availability in summer. 4. Both E. danica and the non‐emitting S. lutaria increased the total N₂O and N₂ emission from sediment in a density‐dependent manner. While N₂O directly emitted by benthic invertebrates can be partially consumed in the sediment (E. danica), non‐emitting species can still indirectly contribute to total N₂O emission from sediment (S. lutaria).     

The fate of assimilated nitrogen in streams: an in situ benthic chamber study

1. Nitrogen (N) processing in streams has been investigated using whole‐stream ¹⁵N addition experiments that, in general, have found that a large proportion of added nitrate removed from the water column appears to be assimilated by the stream benthos. The long‐term fate of this retained N is unknown, and of particular interest is the possibility that it becomes denitrified through coupled mineralisation–nitrification–denitrification processes (indirect denitrification). 2. We used in situ chambers to produce highly ¹⁵N‐enriched benthic biofilms and removed the chambers to allow biofilms to interact with ambient stream conditions. Nitrogen assimilation and direct denitrification were estimated from the first chamber deployment. Chambers were periodically reinstalled over 4 weeks to measure tracer ¹⁵N in ammonium (), nitrate () and dinitrogen (N₂), from which we estimated subsequent rates of biotic N transformations, including N mineralisation (ammonification), nitrification and indirect denitrification. We also estimated rates of depuration of ¹⁵N tracer from benthic biomass compartments. 3. Nitrate uptake was roughly equivalent in the sand and cobble habitats that dominated the stream. Direct denitrification (denitrification of from the water column) was an order of magnitude higher in cobble habitats than in sand habitats, accounting for c. 26 and 2% of total nitrate uptake in cobble and sand, respectively. 4. Mean residence times of actively cycling organic N in stream benthos (algae and microbes) were 16 days in cobble habitats and 9 days in sand habitats. The difference between habitat types was driven by the influence of N residence time in epilithic biofilms (18 days) on cobbles. 5. Release of enriched ¹⁵ was the primary flux of remineralised N, while release of enriched ¹⁵ was an order of magnitude less. We detected slight ¹⁵N enrichment in dissolved nitrogen gas (N₂) in post‐enrichment sampling, indicating that indirect denitrification was taking place. However, indirect denitrification accounted for <0.1% of the assimilated N. 6. These experiments agree with results of whole‐stream ¹⁵N additions, in that most added N was assimilated rather than directly denitrified. Assimilation was primarily a short‐term N retention mechanism in this stream, and indirect denitrification of assimilated N accounted for only a minor proportion of the observed ¹⁵N loss over time. 7. Remaining possible fates include export of N as particulate organic matter, which may lead to additional storage of assimilated N in downstream habitats, and consumption by grazers.     

Changes in grassland ecosystem function due to extreme rainfall events: implications for responses to climate change

Climate change is causing measurable changes in rainfall patterns, and will likely cause increases in extreme rainfall events, with uncertain implications for key processes in ecosystem function and carbon cycling. We examined how variation in rainfall total quantity (Q), the interval between rainfall events (I), and individual event size (S_E) affected soil water content (SWC) and three aspects of ecosystem function: leaf photosynthetic carbon gain (), aboveground net primary productivity (ANPP), and soil respiration (). We utilized rainout shelter‐covered mesocosms (2.6 m³) containing assemblages of tallgrass prairie grasses and forbs. These were hand watered with 16 I×Q treatment combinations, using event sizes from 4 to 53 mm. Increasing Q by 250% (400–1000 mm yr^?1) increased mean soil moisture and all three processes as expected, but only by 20–55% (P≤0.004), suggesting diminishing returns in ecosystem function as Q increased. Increasing I (from 3 to 15 days between rainfall inputs) caused both positive () and negative () changes in ecosystem processes (20–70%, P≤0.01), within and across levels of Q, indicating that I strongly influenced the effects of Q, and shifted the system towards increased net carbon uptake. Variation in S_E at shorter I produced greater response in soil moisture and ecosystem processes than did variation in S_E at longer I, suggesting greater stability in ecosystem function at longer I and a priming effect at shorter I. Significant differences in ANPP and between treatments differing in I and Q but sharing the same S_E showed that the prevailing pattern of rainfall influenced the responses to a given event size. Grassland ecosystem responses to extreme rainfall patterns expected with climate change are, therefore, likely to be variable, depending on how I, Q, and S_E combine, but will likely result in changes in ecosystem carbon cycling.     

Relative influence of bioturbation and predation on organic matter processing in river sediments: a microcosm experiment

1. Our objective was to measure the effects of bioturbation and predation on the physical characteristics and biogeochemical processes in river sediments. 2. We investigated the impacts of tubificid worms tested separately and together with an omnivore (Gammarus pulex), which does feed on tubificids, on sediment distribution, water flux, sediment organic carbon, biofilm biomass and microbial activities, and the concentrations of dissolved oxygen, dissolved organic carbon, PO, NO, NO and NH in slow filtration sand–gravel columns. We hypothesised that gammarids, which exploit the top 2–3 cm of the sediment, would modify the impact of worms at the sediment surface. 3. In experiments both with and without gammarids, bioturbation by the tubificids modified both the distribution of surface particles in the sediment column and water flux. In addition, microbial aerobic (oxygen consumption) and anaerobic (denitrification and fermentative decomposition of organic matter) processes in the sediment were stimulated in the presence of tubificid worms. However, G. pulex did not affect either the density or bioturbation activity of the tubificid worms. 4. Bioturbation by the benthos can be a major process in river habitats, contributing to the retention of organic matter in sediment dynamics. The presence of at least one predator had no effect on bioturbation in sediments. In such systems, physical heterogeneity may be sufficient for tubificids to escape from generalist predators, though more specialised ones might have more effect.     

Periphyton‐based transfer functions to assess ecological imbalance and management of a subtropical ombrotrophic peatland

1. To assess the biological status and response of aquatic resources to management actions, managers and decision‐makers require accurate and precise metrics. This is especially true for some parts of the Florida Everglades where multiple stressors (e.g. hydrologic alterations and eutrophication) have resulted in a highly degraded and fragmented ecosystem. Biological assessments are required that directly allow for the evaluation of historical and current status and responses to implantation of large‐scale restoration projects. 2. Utilising periphyton composition and water‐quality data obtained from long‐term (15 years) monitoring programmes, we developed calibrated and verified periphyton‐based numerical models (transfer functions) that could be used to simultaneously assess multiple stressors affecting the Everglades peatland (e.g. salinity, nitrogen and phosphorus). Periphyton is an ideal indicator because responses to stressors are rapid and predictable and possess valued ecological attributes. 3. Weighted averaging partial least squares regression was used to develop models to infer water‐quality concentrations from 456 samples comprising 319 periphyton taxa. Measured versus periphyton‐inferred concentrations were strongly related for log‐transformed salinity ( = 0.81; RMSEP 0.15 mg L^?1) and log‐transformed total phosphorus (TP; = 0.70; RMSEP 0.18 mg L^?1), but weakly related for total Kjeldahl nitrogen (TKN) ( = 0.46; RMSEP 0.12 mg L^?1). Validation results using an independent 455 sample data set were similar (log(salinity) r² = 0.78, log(TP) r² = 0.65 and log(TKN) r² = 0.38). 4. Water Conservation Area 1 (WCA‐1), a large ombrotrophic subtropical peatland impacted by multiple water‐quality stressors that has undergone major changes in water management, was used as a case study. The models were applied to a long‐term periphyton data set to reconstruct water‐quality trends in relation to restoration efforts to reduce nutrient loading to the Everglades. The combination of biologically inferred TP and salinity was used to identify the ecological status of periphyton assemblages. Periphyton assemblages were ecologically imbalanced with respect to salinity and TP. Salinity imbalance varied spatially and temporally, whereas TP was spatially restricted. Imbalances caused by water management were owing to salinity more so than to TP. 5. The transfer functions developed for the Everglades are trait‐based quantitative numerical methods and are ideal because the abundances of species are modelled numerically in relation to a stressor. The resulting inferred value is a numerical representation of the stressor’s effect on biological condition that can be compared against the management of the stressor independent of other factors. The benefits are that biological lags or hysteresis events can easily be identified and environmental conditions can be estimated when measurements are lacking. Reporting biological assessments in terms of well‐defined water‐quality metrics (e.g. numeric criterion) increased the communicative ability of the assessment. The use of multiple metrics to assess ecological imbalance increased the ability to identify probable causes.