Metabolic regimes of three mid-order streams in southern Ontario,Canada exposed to contrasting sources of nutrients

Light, temperature, and discharge control stream metabolism, but the response of gross primary production (GPP) and ecosystem respiration (ER) to seasonal variation in these physical drivers may differ in accordance with the types of human activities present in the catchment. Our study examined three mid-order streams in southern Ontario, Canada that differed in anthropogenic nutrient sources (i.e., sewage treatment plant effluent, sewage lagoon effluent, and agriculture), but had comparable light, temperature, and discharge regimes. For each stream, GPP and ER were estimated daily from June through November. Comparisons of paired daily metabolic rates revealed pairwise differences among all streams, with streams receiving sewage effluent having greater rates and variability of GPP and ER than the stream draining agricultural land. The two sewage influenced streams differed only in ER. Temporal patterns of GPP and ER were correlated for all streams throughout the study period and were most affected by seasonal variation in temperature, whereby effluent receiving streams responded more rapidly to increases in temperature. Our findings suggest that managers may need to balance effects of human activities with regional environmental constraints on stream metabolism to maintain and enhance the ecological condition and services of stream ecosystems.

     

Multiple Scales of Temporal Variability in Ecosystem Metabolism Rates: Results from 2 Years of Continuous Monitoring in a Forested Headwater Stream

Headwater streams are key sites of nutrient and organic matter processing and retention, but little is known about temporal variability in gross primary production (GPP) and ecosystem respiration (ER) rates as a result of the short duration of most metabolism measurements in lotic ecosystems. We examined temporal variability and controls on ecosystem metabolism by measuring daily rates continuously for 2 years in Walker Branch, a first-order deciduous forest stream. Four important scales of temporal variability in ecosystem metabolism rates were identified: (1) seasonal, (2) day-to-day, (3) episodic (storm-related), and (4) inter-annual. Seasonal patterns were largely controlled by the leaf phenology and productivity of the deciduous riparian forest. Walker Branch was strongly net heterotrophic throughout the year with the exception of the open-canopy spring when GPP and ER rates were co-equal. Day-to-day variability in weather conditions influenced light reaching the streambed, resulting in high day-to-day variability in GPP particularly during spring (daily light levels explained 84% of the variance in daily GPP in April). Episodic storms depressed GPP for several days in spring, but increased GPP in autumn by removing leaves shading the streambed. Storms depressed ER initially, but then stimulated ER to 2–3 times pre-storm levels for several days. Walker Branch was strongly net heterotrophic in both years of the study, with annual GPP being similar (488 and 519 g O₂ m⁻² y⁻¹ or 183 and 195 g C m⁻² y⁻¹) but annual ER being higher in 2004 than 2005 (−1,645 vs. −1,292 g O₂ m⁻² y⁻¹ or −617 and −485 g C m⁻² y⁻¹). Inter-annual variability in ecosystem metabolism (assessed by comparing 2004 and 2005 rates with previous measurements) was the result of the storm frequency and timing and the size of the spring macroalgal bloom. Changes in local climate can have substantial impacts on stream ecosystem metabolism rates and ultimately influence the carbon source and sink properties of these important ecosystems.     

Seasonal weather effects on hydrology drive the metabolism of non-forest lowland streams

Weather variations change stream hydrological conditions, affecting the stream function. A seasonal study in three well-conserved first-order Pampean streams was carried out to test the hypothesis that rainfalls are the main drivers of whole-stream metabolism, through their effects on hydrology. We estimated the stream metabolism and metabolic contribution of six relevant communities (angiosperms, macroalgae, seston, epiphyton, epipelon, and hyporheos) during late spring, summer, and winter and examined the relation between gross primary production (GPP) and photosynthetic active radiation (PAR). Our results showed that the decrease in available streambed light due to the dissolved organic carbon after rainfalls was the main factor related to the decrease in the ecosystem and community metabolisms. For instance, GPP oscillated from ~10 gO₂ m^?2 d^?1 in early spring (low flows) to ~3 gO₂ m^?2 d^?1 in summer (high flows). Ecosystem respiration (ER) was less sensitive than GPP to rainfalls due to the increase of hyporheic respiration. Rainfalls also caused a significant loss of downstream macroalgal biomass. At a day scale, the high PAR of late spring and summer saturated GPP during the afternoon, and the low temperature of winter mornings constrained GPP. Hence, the knowledge of weather changes is key to understanding the main hydrological drivers of stream function.     

Temperature and the metabolic balance of streams

1. It is becoming increasingly clear that fresh waters play a major role in the global C cycle. Stream ecosystem respiration (ER) and gross primary productivity (GPP) exert a significant control on organic carbon fluxes in fluvial networks. However, little is known about how climate change will influence these fluxes. 2. Here, we used a ‘natural experiment’ to demonstrate the role of temperature and nutrient cycling in whole‐system metabolism (ER, GPP and net ecosystem production – NEP), in naturally heated geothermal (5–25 °C) Icelandic streams. 3. We calculated ER and GPP with a new, more accurate method, which enabled us to take into account the additional uncertainties owing to stream spatial heterogeneity in oxygen concentrations within a reach. ER ranged 1–25 g C m^?2 day^?1 and GPP 1–10 g C m^?2 day^?1. The median uncertainties (based on 1 SD) in ER and GPP were 50% and 20%, respectively. 4. Despite extremely low water nutrient concentrations, high metabolic rates in the warm streams were supported by fast cycling rates of nutrients, as revealed from inorganic nutrient (N, P) addition experiments. 5. ER exceeded GPP in all streams (with average GPP/ER = 0.6) and was more strongly related to temperature than GPP, resulting in elevated negative NEP with warming. We show that, as a first approximation based on summer investigations, global stream carbon emission to the atmosphere would nearly double from 0.12 Pg C year^?1 at 13 °C to 0.21 (0.15–0.33) Pg C year^?1 with a 5 °C warming. 6. Compared to previous studies from natural systems (including terrestrial ecosystems), the temperature dependence of stream metabolism was not confounded by latitude or altitude, seasonality, light and nutrient availability, water chemistry, space availability (water transient storage), and water availability. 7. Consequently, stream nutrient processing is likely to increase with warming, protecting downstream ecosystems (rivers, estuaries, coastal marine systems) during the summer low flows from nutrient enrichment, but at the cost of increased CO₂ flux back to the atmosphere.     

Land use controls stream ecosystem metabolism by shifting dissolved organic matter and nutrient regimes

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Flow extremes and benthic organic matter shape the metabolism of a headwater Mediterranean stream

1. Single‐station diel oxygen curves were used to monitor the oxygen metabolism of an intermittent, forested third‐order stream (Fuirosos) in the Mediterranean area, over a period of 22 months. Ecosystem respiration (ER) and gross primary production (GPP) were estimated and related to organic matter inputs and photosynthetically active radiation (PAR) in order to understand the effect of the riparian forest on stream metabolism. 2. Annual ER was 1690 g O₂ m^?2 year^?1 and annual GPP was 275 g O₂ m^?2 year^?1. Fuirosos was therefore a heterotrophic stream, with P : R ratios averaging 0.16. 3. GPP rates were relatively low, ranging from 0.05 to 1.9 g O₂ m^?2 day^?1. The maximum values of GPP occurred during a few weeks in spring, and ended when the riparian canopy was fully closed. The phenology of the riparian vegetation was an important determinant of light availability, and consequently, of GPP. 4. On a daily scale, light and temperature were the most important factors governing the shape of photosynthesis–irradiance (P–I) curves. Several patterns could be generalised in the P–I relationships. Hysteresis‐type curves were characteristic of late autumn and winter. Light saturation responses (that occurred at irradiances higher than 90 μE m^?2 s^?1) were characteristic of early spring. Linear responses occurred during late spring, summer and early autumn when there was no evidence of light saturation. 5. Rates of ER were high when compared with analogous streams, ranging from 0.4 to 32 g O₂ m^?2 day^?1. ER was highest in autumn 2001, when organic matter accumulations on the streambed were extremely high. By contrast, the higher discharge in autumn 2002 prevented these accumulations and caused lower ER. The Mediterranean climate, and in its effect the hydrological regime, were mainly responsible for the temporal variation in benthic organic matter, and consequently of ER.     

Inter‐regional comparison of land‐use effects on stream metabolism

1. Rates of whole‐system metabolism (production and respiration) are fundamental indicators of ecosystem structure and function. Although first‐order, proximal controls are well understood, assessments of the interactions between proximal controls and distal controls, such as land use and geographic region, are lacking. Thus, the influence of land use on stream metabolism across geographic regions is unknown. Further, there is limited understanding of how land use may alter variability in ecosystem metabolism across regions. 2. Stream metabolism was measured in nine streams in each of eight regions (n = 72) across the United States and Puerto Rico. In each region, three streams were selected from a range of three land uses: agriculturally influenced, urban‐influenced, and reference streams. Stream metabolism was estimated from diel changes in dissolved oxygen concentrations in each stream reach with correction for reaeration and groundwater input. 3. Gross primary production (GPP) was highest in regions with little riparian vegetation (sagebrush steppe in Wyoming, desert shrub in Arizona/New Mexico) and lowest in forested regions (North Carolina, Oregon). In contrast, ecosystem respiration (ER) varied both within and among regions. Reference streams had significantly lower rates of GPP than urban or agriculturally influenced streams. 4. GPP was positively correlated with photosynthetically active radiation and autotrophic biomass. Multiple regression models compared using Akaike’s information criterion (AIC) indicated GPP increased with water column ammonium and the fraction of the catchment in urban and reference land‐use categories. Multiple regression models also identified velocity, temperature, nitrate, ammonium, dissolved organic carbon, GPP, coarse benthic organic matter, fine benthic organic matter and the fraction of all land‐use categories in the catchment as regulators of ER. 5. Structural equation modelling indicated significant distal as well as proximal control pathways including a direct effect of land‐use on GPP as well as SRP, DIN, and PAR effects on GPP; GPP effects on autotrophic biomass, organic matter, and ER; and organic matter effects on ER. 6. Overall, consideration of the data separated by land‐use categories showed reduced inter‐regional variability in rates of metabolism, indicating that the influence of agricultural and urban land use can obscure regional differences in stream metabolism.     

Annual cycle and inter-annual variability of gross primary production and ecosystem respiration in a floodprone river during a 15-year period

1. Temporal variation in ecosystem metabolism over a 15‐year period (1986–2000) was evaluated in a seventh order channelised gravel bed river (mean annual discharge 48.7 m³ s^?1) of the Swiss Plateau. The river is subject to frequent disturbance by bed‐moving spates. Daily integrals of gross primary production (GPP) and ecosystem respiration (ER) were calculated based on single‐station diel oxygen curves. 2. Seasonal decomposition of the time series of monthly metabolism rates showed that approximately 50% of the variation of GPP and ER can be attributed to season. Annual GPP averaged 5.0 ± 0.6 g O₂ m^?2 day^?1 and showed no long‐term trend. 3. Ecosystem respiration, averaging 6.2 ± 1.4 g O₂ m^?2 day^?1, declined from 8.8 to 4.1 g O₂ m^?2 day^?1 during the 15‐year period. This significant trend paralleled a decline in nitrate and soluble reactive phosphorus concentrations, and the biochemical oxygen demand discharged by sewage treatment facilities upstream of the study reach. The ratio of GPP to ER (P/R) increased from 0.53 to about 1 as consequence of ER reduction. 4. Bed moving spates reduced GPP by 49% and ER by 19%. Postspate recovery of GPP was rapid between spring and autumn and slow during winter. Recovery of ER lacked any seasonal pattern. Annual patterns of daily GPP and to a minor extent of daily ER can be described as a sequence of recovery periods frequently truncated by spates. 5. The study showed that disturbance by frequent bed‐moving spates resulted in major stochastic variation in GPP and ER but annual patterns were still characterised by a distinct seasonal cycle. It also became evident that stream metabolism is a suitable method to assess effects of gradual changes in water quality.     

The Metabolic Regimes at the Scale of an Entire Stream Network Unveiled Through Sensor Data and Machine Learning

Streams and rivers form dense networks that drain the terrestrial landscape and are relevant for biodiversity dynamics, ecosystem functioning, and transport and transformation of carbon. Yet, resolving in both space and time gross primary production (GPP), ecosystem respiration (ER) and net ecosystem production (NEP) at the scale of entire stream networks has been elusive so far. Here, combining Random Forest (RF) with time series of sensor data in 12 reach sites, we predicted annual regimes of GPP, ER, and NEP in 292 individual stream reaches and disclosed properties emerging from the network they form. We further predicted available light and thermal regimes for the entire network and expanded the library of stream metabolism predictors. We found that the annual network-scale metabolism was heterotrophic yet with a clear peak of autotrophy in spring. In agreement with the River Continuum Concept, small headwaters and larger downstream reaches contributed 16% and 60%, respectively, to the annual network-scale GPP. Our results suggest that ER rather than GPP drives the metabolic stability at the network scale, which is likely attributable to the buffering function of the streambed for ER, while GPP is more susceptible to flow-induced disturbance and fluctuations in light availability. Furthermore, we found large terrestrial subsidies fueling ER, pointing to an unexpectedly high network-scale level of heterotrophy, otherwise masked by simply considering reach-scale NEP estimations. Our machine learning approach sheds new light on the spatiotemporal dynamics of ecosystem metabolism at the network scale, which is a prerequisite to integrate aquatic and terrestrial carbon cycling at relevant scales.

     

Ecosystem metabolism and nutrient uptake in an urban,piped headwater stream

Piped streams, or streams that run underground, are often associated with urbanization. Despite the fact that they are ubiquitous in many urban watersheds, there is little empirical evidence regarding the ecological structure and function of piped stream reaches. This study measured ecosystem metabolism, nutrient uptake, and related characteristics of Pettee Brook—an urban stream that flows through several piped sections in Durham, New Hampshire, USA. Pettee Brook had high chloride and nutrient concentrations, low benthic biomass, and low rates of gross primary productivity (GPP), ecosystem respiration (ER), and nutrient uptake along its entire length during summer. Spring was a period of elevated biological activity, as increased light availability in the un-piped sections of the stream led to substantially higher GPP, ER, NH₄ uptake, and PO₄ uptake in these open reaches. Piped reaches of Pettee Brook were similar to open reaches in terms of water quality, dissolved O₂ concentration, temperature, and discharge. Piped reaches did, however, have significantly less light, shallower sediments, and no debris dams. The absence of light inhibited autotrophic activity in piped reaches, resulting in the complete loss of GPP as well as a significant reduction in benthic AFDM and chlorophyll a biomass. Heterotrophic activity in piped reaches was not impaired to the same extent as autotrophic activity. Reduced ER was observed in piped reaches during the summer, but we failed to find significantly lower DOC or nutrient uptake rates in piped reaches than in open reaches. Carbon consumption in piped reaches, which do not have significant autochthonous or allochthonous carbon replenishment, must rely primarily on upstream inputs of organic matter. These results suggest that although ecological conditions in piped streams may be degraded beyond the extent of other urban stream reaches, piped reaches may still sustain some measurable ecosystem function.     

An approach for the identification of exemplar sites for scaling up targeted field observations of benthic biogeochemistry in heterogeneous environments

Headwater streams influence the biogeochemical characteristics of large rivers and play important roles in regional and global carbon budgets. The combined effects of seasonality and land use change on the biogeochemistry of headwater streams, however, are not well understood. In this study we assessed the influence of catchment land use and seasonality on the composition of dissolved organic matter (DOM) and ecosystem metabolism in headwater streams of a Kenyan river. Fifty sites in 34 streams draining a gradient of catchment land use from 100% natural forest to 100% agriculture were sampled to determine temporal and spatial variation in DOM composition. Gross primary production (GPP) and ecosystem respiration (ER) were determined in 10 streams draining primarily forest or agricultural catchments. Absorbance and fluorescence spectrophotometry of DOM reflected notable shifts in composition along the land use gradient and with season. During the dry season, forest streams contained higher molecular weight and terrestrially derived DOM, whereas agricultural streams were dominated by autochthonous production and low molecular weight DOM. During the rainy season, aromaticity and high molecular weight DOM increased in agricultural streams, coinciding with seasonal erosion of soils and inputs of organic matter from farmlands. Most of the streams were heterotrophic. However, GPP and ER were generally greater in agricultural streams, driven by higher dissolved nutrient (mainly TDN) concentrations, light availability (open canopy) and temperature compared with forest streams. There were correlations between freshly and autochthonously produced DOM, GPP and ER during both the dry and wet seasons. This is one of the few studies to link land-use with organic carbon dynamics and DOM composition. Measures of ecosystem metabolism in these streams help to affirm the role of tropical streams and rivers as important components of the global carbon cycle and demonstrate that even semi-intensive, smallholder agriculture can have measurable effects on riverine ecosystem functioning.     

Climate-Induced Changes in Spring Snowmelt Impact Ecosystem Metabolism and Carbon Fluxes in an Alpine Stream Network

Although stream ecosystems are recognized as an important component of the global carbon cycle, the impacts of climate-induced hydrological extremes on carbon fluxes in stream networks remain unclear. Using continuous measurements of ecosystem metabolism, we report on the effects of changes in snowmelt hydrology during the anomalously warm winter 2013/2014 on gross primary production (GPP), ecosystem respiration (ER), and net ecosystem production (NEP) in an Alpine stream network. We estimated ecosystem metabolism across 12 study reaches of the 254 km² subalpine Ybbs River Network (YRN), Austria, for 18 months. During spring snowmelt, GPP peaked in 10 of our 12 study reaches, which appeared to be driven by PAR and catchment area. In contrast, the winter precipitation shift from snow to rain following the low-snow winter in 2013/2014 increased spring ER in upper elevation catchments, causing spring NEP to shift from autotrophy to heterotrophy. Our findings suggest that the YRN transitioned from a transient sink to a source of carbon dioxide (CO₂) in spring as snowmelt hydrology differed following the high-snow versus low-snow winter. This shift toward increased heterotrophy during spring snowmelt following a warm winter has potential consequences for annual ecosystem metabolism, as spring GPP contributed on average 33% to annual GPP fluxes compared to spring ER, which averaged 21% of annual ER fluxes. We propose that Alpine headwaters will emit more within-stream respiratory CO₂ to the atmosphere while providing less autochthonous organic energy to downstream ecosystems as the climate gets warmer.     

The influence of native replanting on stream ecosystem metabolism in a degraded landscape: can a little vegetation go a long way?

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Inter-biome comparison of factors controlling stream metabolism

1. We studied whole-ecosystem metabolism in eight streams from several biomes in North America to identify controls on the rate of stream metabolism over a large geographic range. The streams studied had climates ranging from tropical to cool-temperate and from humid to arid and were all relatively uninfluenced by human disturbances.
2. Rates of gross primary production (GPP), ecosystem respiration (R) and net ecosystem production (NEP) were determined using the open-system, two-station diurnal oxygen change method.
3. Three general patterns in metabolism were evident among streams: (1) relatively high GPP with positive NEP (i.e. net oxygen production) in early afternoon, (2) moderate primary production with a distinct peak in GPP during daylight but negative NEP at all times and (3) little or no evidence of GPP during daylight and a relatively constant and negative NEP over the entire day.
4. Gross primary production was most strongly correlated with photosynthetically active radiation (PAR). A multiple regression model that included log PAR and stream water soluble reactive phosphorus (SRP) concentration explained 90% of the variation in log GPP.
5. Ecosystem respiration was significantly correlated with SRP concentration and size of the transient storage zone and, together, these factors explained 73% of the variation in R. The rate of R was poorly correlated with the rate of GPP.
6. Net ecosystem production was significantly correlated only with PAR, with 53% of the variation in log NEP explained by log PAR. Only Sycamore Creek, a desert stream in Arizona, had positive NEP (GPP: R  > 1), supporting the idea that streams are generally net sinks rather than net sources of organic matter.
7. Our results suggest that light, phosphorus concentration and channel hydraulics are important controls on the rate of ecosystem metabolism in streams over very extensive geographic areas.     

Watershed-scale Land Use Change Increases Ecosystem Metabolism in an Agricultural Stream

Ecosystems - Stream metabolism, in the form of gross primary production (GPP) and ecosystem respiration (ER), is an important metric of stream ecosystem function, given GPP and ER are integrative...     

Multi‐scaled drivers of rural prairie stream metabolism along human activity gradients

1. Stream metabolism is increasingly used for monitoring and assessment of the biological condition of aquatic ecosystems. However, distal environmental drivers, such as land use, are typically not well connected to the proximate controls, such as stream chemistry, that are usually invoked as driving metabolism. This is particularly true for North American prairie streams and for grassland streams worldwide. 2. Stream metabolism was measured at the outflow of 19 subcatchments of the Red River in southern Manitoba, Canada. Subcatchments represented gradients of nutrient‐producing human activities present in the region, that is, wastewater treatment (WWT), livestock production and crop cultivation. Stream metabolism was estimated at all sites using diel changes in dissolved oxygen (DO) concentration over 1 week in the middle of summer. Environmental parameters hypothesised to control stream metabolism were sampled across three spatial scales (stream reach, stream segment and catchment). Model selection using Akaike’s information criterion (AIC) was used to determine linkages between environmental parameters and measures of stream metabolism. 3. Estimated rates of metabolism were within the range of past studies of metabolism in prairie streams, although most streams had negative values of net ecosystem metabolism. However, production‐to‐respiration ratios were >0.5, at all but three sites suggesting that autochthonous production was an important source of organic matter. 4. The a priori model that best predicted gross primary production (GPP) was the intensity of nutrient‐producing human activities (i.e. WWT, livestock and crop cultivation) measured at the catchment scale. Ecosystem respiration (ER) was best predicted by the a priori model comprised of GPP, total nitrogen (TN) and total phosphorus (TP). However, model averaging revealed that prediction of ER could be improved by including riparian cover and removing TP from the model. The positive association between GPP and ER suggested that heterotrophic compartments of the ecosystem were modest contributors to variation in respiration rates. 5. Overall, this study suggests that variation of metabolism in prairie streams of southern Manitoba is controlled by human activities occurring at the catchment scale, a finding consistent with current hierarchically structured riverine paradigms. Moreover, increased understanding of the hierarchical structure of stream metabolism drivers will help to ensure that assessment results can be used more effectively to inform management strategies for prairie ecosystems.     

Metabolism,Gas Exchange,and Carbon Spiraling in Rivers

Ecosystem metabolism, that is, gross primary productivity (GPP) and ecosystem respiration (ER), controls organic carbon (OC) cycling in stream and river networks and is expected to vary predictably with network position. However, estimates of metabolism in small streams outnumber those from rivers such that there are limited empirical data comparing metabolism across a range of stream and river sizes. We measured metabolism in 14 rivers (discharge range 14–84 m³ s^?1) in the Western and Midwestern United States (US). We estimated GPP, ER, and gas exchange rates using a Lagrangian, 2-station oxygen model solved in a Bayesian framework. GPP ranged from 0.6–22 g O₂ m^?2 d^?1 and ER tracked GPP, suggesting that autotrophic production supports much of riverine ER in summer. Net ecosystem production, the balance between GPP and ER was 0 or greater in 4 rivers showing autotrophy on that day. River velocity and slope predicted gas exchange estimates from these 14 rivers in agreement with empirical models. Carbon turnover lengths (that is, the distance traveled before OC is mineralized to CO₂) ranged from 38 to 1190 km, with the longest turnover lengths in high-sediment, arid-land rivers. We also compared estimated turnover lengths with the relative length of the river segment between major tributaries or lakes; the mean ratio of carbon turnover length to river length was 1.6, demonstrating that rivers can mineralize much of the OC load along their length at baseflow. Carbon mineralization velocities ranged from 0.05 to 0.81 m d^?1, and were not different than measurements from small streams. Given high GPP relative to ER, combined with generally short OC spiraling lengths, rivers can be highly reactive with regard to OC cycling.     

Resistance and resilience of ecosystem metabolism in a flood-prone river system

1. Gross primary production (GPP) and ecosystem respiration (ER) were analysed for 18 months in two reaches of the River Thur, a prealpine river in Switzerland. The upper reach at 655 m above sea level (a.s.l.) is bedrock constrained, has a high slope (0.60%) and a catchment area of 126 km2. The lower reach at 370 m a.s.l. has a more extensive hyporheic zone, a lower slope (0.17%) and a catchment of 1696 km2.
2. In both reaches, temporal patterns of stream metabolism reflected the occurrence of bed-moving spates. Average reductions of GPP and ER by spates were 53 and 24% in the upper reach, and 37 and 14% in the lower reach, respectively. The greater resistance of ER than GPP in both reaches shifted the ecosystem metabolism towards heterotrophy (decrease of the ratio of GPP to ER (P/R)) following spates.
3. Recovery of GPP was significantly faster in the lower reach and exhibited distinct seasonal variation (positive correlation with incident light). The differences in stability (both resistance and resilience) between reaches reflected differences in geomorphic settings and disturbance regime.
4. Stepwise regression analysis was used to explore the potential influence of season, disturbance and prevailing environmental conditions on stream metabolism in each reach. Time since spate plus temperature explained 73 and 86% of variation in ER and GPP, respectively, in the upper reach and 55% of variation in ER in the lower reach. Season plus prevailing environmental conditions explained 67% of variation in GPP in the lower reach.
5. To test how the perception of stability may change with increasing scale of observation, the disturbance regimes of 12 sites were compared with the disturbance regime of the entire Thur catchment. The analysis suggests that stream metabolism at the catchment scale is far more resistant to high flow events than at the reach scale.     

Seasonal hysteresis of net ecosystem exchange in response to temperature change: patterns and causes

Understanding how net ecosystem exchange (NEE) changes with temperature is central to the debate on climate change‐carbon cycle feedbacks, but still remains unclear. Here, we used eddy covariance measurements of NEE from 20 FLUXNET sites (203 site‐years of data) in mid‐ and high‐latitude forests to investigate the temperature response of NEE. Years were divided into two half thermal years (increasing temperature in spring and decreasing temperature in autumn) using the maximum daily mean temperature. We observed a parabolic‐like pattern of NEE in response to temperature change in both the spring and autumn half thermal years. However, at similar temperatures, NEE was considerably depressed during the decreasing temperature season as compared with the increasing temperature season, inducing a counter‐clockwise hysteresis pattern in the NEE–temperature relation at most sites. The magnitude of this hysteresis was attributable mostly (68%) to gross primary production (GPP) differences but little (8%) to ecosystem respiration (ER) differences between the two half thermal years. The main environmental factors contributing to the hysteresis responses of NEE and GPP were daily accumulated radiation. Soil water content (SWC) also contributed to the hysteresis response of GPP but only at some sites. Shorter day length, lower light intensity, lower SWC and reduced photosynthetic capacity may all have contributed to the depressed GPP and net carbon uptake during the decreasing temperature seasons. The resultant hysteresis loop is an important indicator of the existence of limiting factors. As such, the role of radiation, LAI and SWC should be considered when modeling the dynamics of carbon cycling in response to temperature change.     

Effects of urban stream burial on nitrogen uptake and ecosystem metabolism: implications for watershed nitrogen and carbon fluxes

Urbanization has resulted in the extensive burial and channelization of headwater streams, yet little is known about the impacts of stream burial on ecosystem functions critical for reducing downstream nitrogen (N) and carbon (C) exports. In order to characterize the biogeochemical effects of stream burial on N and C, we measured NO₃ ^? uptake (using ¹⁵N-NO₃ ^? isotope tracer releases) and gross primary productivity (GPP) and ecosystem respiration (ER) (using whole stream metabolism measurements). Experiments were carried out during four seasons, in three paired buried and open stream reaches, within the Baltimore Ecosystem Study Long-term Ecological Research site. Stream burial increased NO₃ ^? uptake lengths by a factor of 7.5 (p < 0.01) and decreased NO₃ ^? uptake velocity and areal NO₃ ^? uptake rate by factors of 8.2 (p < 0.05) and 9.6 (p < 0.001), respectively. Stream burial decreased GPP by a factor of 11.0 (p < 0.01) and decreased ER by a factor of 5.0 (p < 0.05). From fluorescence Excitation Emissions Matrices analysis, buried streams were found to have significantly altered C quality, showing less labile dissolved organic matter. Furthermore, buried streams had significantly lower transient storage (TS) and water temperatures. Differences in NO₃ ^? uptake, GPP, and ER in buried streams, were primarily explained by decreased TS, light availability, and C quality, respectively. At the watershed scale, we estimate that stream burial decreases NO₃ ^? uptake by 39 % and C production by 194 %. Overall, our results suggest that stream burial significantly impacts NO₃ ^? uptake, stream metabolism, and the quality of organic C exported from watersheds. Given the large impacts of stream burial on stream ecosystem processes, daylighting or de-channelization of streams, through hydrologic floodplain reconnection, may have the potential to alter ecosystem functions in urban watersheds, when used appropriately.