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.     

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.     

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.     

Impact of seasonal changes in stream metabolism on nitrate concentrations in an urban stream

Nitrate (NO₃ ^?) dynamics in urban streams differ from many natural streams due to stormwater runoff, sewage inputs, decreased groundwater discharge, often limited hyporheic exchange, increased primary productivity, and limited carbon input. We investigated NO₃ ^? dynamics in a first-order urban stream in Syracuse, NY, which has urbanized headwaters and a geomorphologically natural downstream section. Twice-monthly water sampling, NO₃ ^? injection tests, NO₃ ^? isotopic analysis, filamentous algae mat density, and riparian shading were used to identify processes regulating NO₃ ^? dynamics in the stream over a 12-month period. The urban headwater reach had low NO₃ ^? (0.006–0.2 mg N/L) in the spring through fall, with a minimum uptake length of 900 m, no canopy cover, and high algae mat density. The downstream natural reach (100% canopy cover during the summer and low algae mat density) had nitrate concentrations between 0.6 and 1.2 mg N/L from winter to summer, which decreased during autumn leaf-off. In the urban reach, autotrophic uptake by filamentous green algae is a major NO₃ ^? sink in summer. In the natural reach, the addition of organic matter to the stream at leaf-off led to a decrease in NO₃ ^? concentration followed by an increase in NO₃ ^? concentration in winter as gross primary productivity decreased. This study shows that the balance between autotrophy and heterotrophy in urban streams is variable and depends on an interplay of drivers such as temperature, light, and carbon inputs that are mediated by the riparian ecosystem.     

Carbon sequestration in a high-elevation, subalpine forest

We studied net ecosystem CO₂ exchange (NEE) dynamics in a high‐elevation, subalpine forest in Colorado, USA, over a two‐year period. Annual carbon sequestration for the forest was 6.71 mol C m^?2 (80.5 g C m^?2) for the year between November 1, 1998 and October 31, 1999, and 4.80 mol C m^?2 (57.6 g C m^?2) for the year between November 1, 1999 and October 31, 2000. Despite its evergreen nature, the forest did not exhibit net CO₂ uptake during the winter, even during periods of favourable weather. The largest fraction of annual carbon sequestration occurred in the early growing‐season; during the first 30 days of both years. Reductions in the rate of carbon sequestration after the first 30 days were due to higher ecosystem respiration rates when mid‐summer moisture was adequate (as in the first year of the study) or lower mid‐day photosynthesis rates when mid‐summer moisture was not adequate (as in the second year of the study). The lower annual rate of carbon sequestration during the second year of the study was due to lower rates of CO₂ uptake during both the first 30 days of the growing season and the mid‐summer months. The reduction in CO₂ uptake during the first 30 days of the second year was due to an earlier‐than‐normal spring warm‐up, which caused snow melt during a period when air temperatures were lower and atmospheric vapour pressure deficits were higher, compared to the first 30 days of the first year. The reduction in CO₂ uptake during the mid‐summer of the second year was due to an extended drought, which was accompanied by reduced latent heat exchange and increased sensible heat exchange. Day‐to‐day variation in the daily integrated NEE during the summers of both years was high, and was correlated with frequent convective storm clouds and concomitant variation in the photosynthetic photon flux density (PPFD). Carbon sequestration rates were highest when some cloud cover was present, which tended to diffuse the photosynthetic photon flux, compared to periods with completely clear weather. The results of this study are in contrast to those of other studies that have reported increased annual NEE during years with earlier‐than‐normal spring warming. In the current study, the lower annual NEE during 2000, the year with the earlier spring warm‐up, was due to (1) coupling of the highest seasonal rates of carbon sequestration to the spring climate, rather than the summer climate as in other forest ecosystems that have been studied, and (2) delivery of snow melt water to the soil when the spring climate was cooler and the atmosphere drier than in years with a later spring warm‐up. Furthermore, the strong influence of mid‐summer precipitation on CO₂ uptake rates make it clear that water supplied by the spring snow melt is a seasonally limited resource, and summer rains are critical for sustaining high rates of annual carbon sequestration.     

Effects of seasonal and interannual variations in leaf photosynthesis and canopy leaf area index on gross primary production of a cool-temperate deciduous broadleaf forest in Takayama, Japan

Revealing the seasonal and interannual variations in forest canopy photosynthesis is a critical issue in understanding the ecological mechanisms underlying the dynamics of carbon dioxide exchange between the atmosphere and deciduous forests. This study examined the effects of temporal variations of canopy leaf area index (LAI) and leaf photosynthetic capacity [the maximum velocity of carboxylation (V _cmax)] on gross primary production (GPP) of a cool-temperate deciduous broadleaf forest for 5 years in Takayama AsiaFlux site, central Japan. We made two estimations to examine the effects of canopy properties on GPP; one is to incorporate the in situ observation of V _cmax and LAI throughout the growing season, and another considers seasonality of LAI but constantly high V _cmax. The simulations indicated that variation in V _cmax and LAI, especially in the leaf expansion period, had remarkable effects on GPP, and if V _cmax was assumed constant GPP will be overestimated by 15%. Monthly examination of air temperature, radiation, LAI and GPP suggested that spring temperature could affect canopy phenology, and also that GPP in summer was determined mainly by incoming radiation. However, the consequences among these factors responsible for interannual changes of GPP are not straightforward since leaf expansion and senescence patterns and summer meteorological conditions influence GPP independently. This simulation based on in situ ecophysiological research suggests the importance of intensive consideration and understanding of the phenology of leaf photosynthetic capacity and LAI to analyze and predict carbon fixation in forest ecosystems.     

Delayed herbivory by migratory geese increases summer‐long CO2 uptake in coastal western Alaska

The advancement of spring and the differential ability of organisms to respond to changes in plant phenology may lead to “phenological mismatches” as a result of climate change. One potential for considerable mismatch is between migratory birds and food availability in northern breeding ranges, and these mismatches may have consequences for ecosystem function. We conducted a three‐year experiment to examine the consequences for CO₂ exchange of advanced spring green‐up and altered timing of grazing by migratory Pacific black brant in a coastal wetland in western Alaska. Experimental treatments represent the variation in green‐up and timing of peak grazing intensity that currently exists in the system. Delayed grazing resulted in greater net ecosystem exchange (NEE) and gross primary productivity (GPP), while early grazing reduced CO₂ uptake with the potential of causing net ecosystem carbon (C) loss in late spring and early summer. Conversely, advancing the growing season only influenced ecosystem respiration (ER), resulting in a small increase in ER with no concomitant impact on GPP or NEE. The experimental treatment that represents the most likely future, with green‐up advancing more rapidly than arrival of migratory geese, results in NEE changing by 1.2 µmol m^?2 s^?1 toward a greater CO₂ sink in spring and summer. Increased sink strength, however, may be mitigated by early arrival of migratory geese, which would reduce CO₂ uptake. Importantly, while the direct effect of climate warming on phenology of green‐up has a minimal influence on NEE, the indirect effect of climate warming manifest through changes in the timing of peak grazing can have a significant impact on C balance in northern coastal wetlands. Furthermore, processes influencing the timing of goose migration in the winter range can significantly influence ecosystem function in summer habitats.     

Seasonal and annual variation of carbon exchange in an evergreen Mediterranean forest in southern France

We present 9 years of eddy covariance measurements made over an evergreen Mediterranean forest in southern France. The goal of this study was to quantify the different components of the carbon (C) cycle, gross primary production (GPP) and ecosystem respiration (R_eco), and to assess the effects of climatic variables on these fluxes and on the net ecosystem exchange of carbon dioxide. The Puéchabon forest acted as a net C sink of ?254 g C m^?2 yr^?1, with a GPP of 1275 g C m^?2 yr^?1 and a R_eco of 1021 g C m^?2 yr^?1. On average, 83% of the net annual C sink occurred between March and June. The effects of exceptional events such the insect‐induced partial canopy defoliation that occurred in spring 2005, and the spring droughts of 2005 and 2006 are discussed. A high interannual variability of ecosystem C fluxes during summer and autumn was observed but the resulting effect on the annual net C budget was moderate. Increased severity and/or duration of summer drought under climate change do not appear to have the potential to negatively impact the average C budget of this ecosystem. On the contrary, factors affecting ecosystem functioning (drought and/or defoliation) during March–June period may reduce dramatically the annual C balance of evergreen Mediterranean forests.     

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.     

Nitrate removal in two relict oxbow urban wetlands: a 15N mass-balance approach

A ¹⁵N-tracer method was used to quantify nitrogen (N) removal processes in two relict oxbow wetlands located adjacent to the Minebank Run restored stream reach in Baltimore County (Maryland, USA) during summer 2009 and early spring 2010. A mass-balance approach was used to directly determine the flow of ¹⁵NO₃ ^? to plants, algae, and sediments, with unaccounted for ¹⁵N assumed to be denitrified. During the summer, plant and algal uptake accounted for 42%, of the added ¹⁵NO₃ ^? in oxbow 1, less than 1% remained in the water column and 57% was unaccounted for. In oxbow 2 during the summer, plant and algal uptake accounted for 63% of the added ¹⁵NO₃ ^?, with <1% remaining in the water column and 38% unaccounted for. During the early spring, plant and algal uptake were much lower in both oxbows, ranging from 0.05 to 13.3% of the ¹⁵N added, with 97 and 87% was unaccounted for in oxbow 1 and 2, respectively. The amount of unaccounted for ¹⁵N was equivalent to estimated areal denitrification rates of 12 and 6 mg N m^?2 d^?1 in the summer and 78 and 15 mg N m^?2 d^?1 in the spring, in oxbow 1 and oxbow 2, respectively. However, the uncertainty of these estimates is high as it was difficult to detect accumulation of ¹⁵N in the sediments which could have accounted for a very large percentage of the added ¹⁵N. Our results suggest that the two relict oxbow wetlands are sinks for NO₃ ^? during both summer and spring but that the pathways of removal vary with plants and algae playing a major role in summer but not in spring.     

Direct and indirect climate change effects on carbon dioxide fluxes in a thawing boreal forest–wetland landscape

In the sporadic permafrost zone of northwestern Canada, boreal forest carbon dioxide (CO₂) fluxes will be altered directly by climate change through changing meteorological forcing and indirectly through changes in landscape functioning associated with thaw‐induced collapse‐scar bog (‘wetland’) expansion. However, their combined effect on landscape‐scale net ecosystem CO₂ exchange (NEE_LAND), resulting from changing gross primary productivity (GPP) and ecosystem respiration (ER), remains unknown. Here, we quantify indirect land cover change impacts on NEE_LAND and direct climate change impacts on modeled temperature‐ and light‐limited NEE_LAND of a boreal forest–wetland landscape. Using nested eddy covariance flux towers, we find both GPP and ER to be larger at the landscape compared to the wetland level. However, annual NEE_LAND (?20 g C m^?2) and wetland NEE (?24 g C m^?2) were similar, suggesting negligible wetland expansion effects on NEE_LAND. In contrast, we find non‐negligible direct climate change impacts when modeling NEE_LAND using projected air temperature and incoming shortwave radiation. At the end of the 21st century, modeled GPP mainly increases in spring and fall due to reduced temperature limitation, but becomes more frequently light‐limited in fall. In a warmer climate, ER increases year‐round in the absence of moisture stress resulting in net CO₂ uptake increases in the shoulder seasons and decreases during the summer. Annually, landscape net CO₂ uptake is projected to decline by 25 ± 14 g C m^?2 for a moderate and 103 ± 38 g C m^?2 for a high warming scenario, potentially reversing recently observed positive net CO₂ uptake trends across the boreal biome. Thus, even without moisture stress, net CO₂ uptake of boreal forest–wetland landscapes may decline, and ultimately, these landscapes may turn into net CO₂ sources under continued anthropogenic CO₂ emissions. We conclude that NEE_LAND changes are more likely to be driven by direct climate change rather than by indirect land cover change impacts.     

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.     

Soil and stream water acidification in a forested catchment in central Japan

Rapid industrialization in East Asia is causing adverse effects due to atmospheric deposition in terrestrial and freshwater ecosystems. Decreasing stream pH and alkalinity and increasing NO₃ ^? concentrations were observed throughout the 1990s in the forested Lake Ijira catchment in central Japan. We investigated these changes using data on atmospheric deposition, soil chemistry, stream water chemistry, and forest growth. Average atmospheric depositions (wet + dry) of 0.83, 0.57, and 1.37 kmol ha^?1 year^?1 for hydrogen, sulfur, and nitrogen, respectively, were among the highest levels in Japan. Atmospheric deposition generally decreased before 1994 and increased thereafter. The catchment was acid-sensitive; stream alkalinity was low (134 μmol_c l^?1) and pH in surface mineral soils decreased from 4.5 in 1990 to 3.9 in 2003. Stream NO₃ ^? concentration nearly doubled (from 22 to 42 μmol_c l^?1) from the late 1980s to the early 2000s. Stream NO₃ ^? concentration was controlled primarily by water temperature before 1996/1997 and by stream discharge thereafter. Stream NO₃ ^? concentrations decreased during the growing season before 1996/1997, but this seasonality was lost thereafter. The catchment became nitrogen-saturated (changing from stage 1 to 2) in 1996/1997, possibly because of declining forest growth rates due to the 1994 summer drought, defoliation of Japanese red pine by pine wilt disease, maturation of Japanese cedar stands, and stimulation of nitrogen mineralization and nitrification due to alkalinization of soils (increased exchangeable Ca²⁺ and soil pH) after the summer drought. Stream pH and alkalinity began decreasing in 1996/1997. The enhanced growing-season NO₃ ^? discharge since 1996/1997 appeared to be the major cause of stream acidification. Increased atmospheric deposition since 1994 may have contributed to this change.     

Exceptional carbon uptake in European forests during the warm spring of 2007: a data–model analysis

Temperate and boreal forests undergo drastic functional changes in the springtime, shifting within a few weeks from net carbon (C) sources to net C sinks. Most of these changes are mediated by temperature. The autumn 2006–winter 2007 record warm period was followed by an exceptionally warm spring in Europe, making spring 2007 a good candidate for advances in the onset of the photosynthetically active period. An analysis of a decade of eddy covariance data from six European forests stands, which encompass a wide range of functional types (broadleaf evergreen, broadleaf deciduous, needleleaf evergreen) and a wide latitudinal band (from 44° to 62°N), revealed exceptional fluxes during spring 2007. Gross primary productivity (GPP) of spring 2007 was the maximum recorded in the decade examined for all sites but a Mediterranean evergreen forest (with a +40 to +130 gC m^?2 anomaly compared with the decadal mean over the January–May period). Total ecosystem respiration (TER) was also promoted during spring 2007, though less anomalous than GPP (with a +17 to +93 gC m^?2 anomaly over 5 months), leading to higher net uptake than the long‐term mean at all sites (+12 to +79 gC m^?2 anomaly over 5 months). A correlative analysis relating springtime C fluxes to simple phenological indices suggested spring C uptake and temperatures to be related. The CASTANEA process‐based model was used to disentangle the seasonality of climatic drivers (incoming radiation, air and soil temperatures) and biological drivers (canopy dynamics, thermal acclimation of photosynthesis to low temperatures) on spring C fluxes along the latitudinal gradient. A sensitivity analysis of model simulations evidenced the roles of (i) an exceptional early budburst combined with elevated air temperature in deciduous sites, and (ii) an early relief of winter thermal acclimation in coniferous sites for the promotion of 2007 spring assimilation.     

Macrophytic, epipelic and epilithic primary production in a semiarid Mediterranean stream

Summary 1. Primary production by Chara vulgaris and by epipelic and epilithic algal assemblages was measured in a semiarid, Mediterranean stream (Chicamo stream, Murcia, Spain) during one annual cycle. 2. The rates of gross primary production (GPP) and community respiration (CR) were determined for each algal assemblage using oxygen change in chambers. The net daily metabolism (NDM) and the GPPd^?1 : CR₂₄ ratio were estimated by patch‐weighting the assemblage‐level metabolism values. 3. Gross primary production and CR showed significant differences between assemblages and dates. The highest rates were measured in summer and spring, while December was the only month when there were no significant differences in either parameters between assemblages. GPP was strongly correlated with respiration, but not with algal biomass. 4. Chara vulgaris showed the highest mean annual metabolic rates (GPP = 2.80 ± 0.83 gC m^?2 h^?1, CR = 0.76 ± 0.29 gC m^?2 h^?1), followed by the epilithic assemblage (GPP = 1.97 ± 0.73 gC m^?2 h^?1, CR = 0.41 ± 0.12 gC m^?2 h^?1) and epipelic algae (GPP = 1.36 ± 0.22 gC m^?2 h^?1, CR = 0.39 ± 0.06 gC m^?2 h^?1). 5. The epipelic assemblage dominated in terms of biomass (82%) and areal cover (88%), compared with the other primary producers. Epipelic algae contributed 84% of gross primary production and 86% of community respiration in the stream. 6. Mean monthly air temperature was the best single predictor of macrophyte respiration and of epipelic GPP and CR. However, ammonium concentration was the best single predictor of C. vulgaris GPP, and suspended solid concentration of epilithon GPP and CR. 7. Around 70% of the variation in both mean GPP and mean CR was explained by the mean monthly air temperature alone. A multiple regression model that included conductivity, PAR and nitrates in addition to mean monthly air temperature, explained 99.99% of the variation in mean CR. 8. Throughout the year, NDM was positive (mean value 7.03 gC m^?2 day^?1), while the GPP : CR₂₄ ratio was higher than 1, confirming the net autotrophy of the system.     

Seasonal controls on interannual variability in carbon dioxide exchange of a near-end-of rotation Douglas-fir stand in the Pacific Northwest, 1997–2006

This study analyzes 9 years of eddy‐covariance (EC) data carried out in a Pacific Northwest Douglas‐fir (Pseudotsuga menzesii) forest (58‐year old in 2007) on the east coast of Vancouver Island, Canada, and characterizes the seasonal and interannual variability in net ecosystem productivity (NEP), gross primary productivity (GPP), and ecosystem respiration (R_e) and primary climatic controls on these fluxes. The annual values (± SD) of NEP, GPP and R_e were 357 ± 51, 2124 ± 125, and 1767 ± 146 g C m^?2 yr^?1, respectively, with ranges of 267–410, 1592–2338, and 1642–2071 g C m^?2 yr^?1, respectively. Spring to early summer (March–June) accounted for more than 80% of annual NEP while late spring to early autumn (May–August) was mainly responsible for its interannual variability (～80%). The major drivers of interannual variability in annual carbon (C) fluxes were annual and spring mean air temperatures (T_a) and water deficiency during late summer and autumn (July–October) when this Douglas‐fir forest growth was often water‐limited. Photosynthetically active radiation (Q), and the combination of Q and soil water content (θ) explained 85% and 91% of the variance of monthly GPP, respectively; and 91% and 96% of the variance of monthly R_e was explained by T_a and the combination of T_a and θ, respectively. Annual net C sequestration was high during optimally warm and normal precipitation years, but low in unusually warm or severely dry years. Excluding 1998 and 1999, the 2 years strongly affected by an El Niño/La Niña cycle, annual NEP significantly decreased with increasing annual mean T_a. Annual NEP will likely decrease whereas both annual GPP and R_e will likely increase if the future climate at the site follows a trend similar to that of the past 40 years.     

Differential responses of root uptake kinetics of NH4+ and NO3− to enriched atmospheric CO2 concentration in field-grown loblolly pine

The nitrogen requirement of plants is predominantly supplied by NH₄⁺ and/or NO₃^? from the soil solution, but the energetic cost of uptake and assimilation is generally higher for NO₃^? than for NH₄⁺. We found that CO₂ enrichment of the atmosphere enhanced the root uptake capacity for NO₃^?, but not for NH₄⁺, in field-grown loblolly pine saplings. Increased preference for NO₃^? at the elevated CO₂ concentration was accompanied by increased carbohydrate levels in roots. The results have important implications for the potential consequences of global climate change on plant-and ecosystem-level processes in many temperate forest ecosystems.     

Effects of urban stream burial on organic matter dynamics and reach scale nitrate retention

Nitrogen (N) retention in streams is an important ecosystem service that may be affected by the widespread burial of streams in stormwater pipes in urban watersheds. We predicted that stream burial suppresses the capacity of streams to retain nitrate (NO₃ ^?) by eliminating primary production, reducing respiration rates and organic matter availability, and increasing specific discharge. We tested these predictions by measuring whole-stream NO₃ ^? removal rates using ¹⁵NO₃ ^? isotope tracer releases in paired buried and open reaches in three streams in Cincinnati, Ohio (USA) during four seasons. Nitrate uptake lengths were 29 times greater in buried than open reaches, indicating that buried reaches were less effective at retaining NO₃ ^? than open reaches. Burial suppressed NO₃ ^? retention through a combination of hydrological and biological processes. The channel shape of two of the buried reaches increased specific discharge which enhanced NO₃ ^? transport from the channel, highlighting the relationship between urban infrastructure and ecosystem function. Uptake lengths in the buried reaches were further lengthened by low stream biological NO₃ ^? demand, as indicated by NO₃ ^? uptake velocities 17-fold lower than that of the open reaches. We also observed differences in the periphyton enzyme activity between reaches, indicating that the effects of burial cascade from the microbial to the ecosystem scale. Our results suggest that stream restoration practices involving “daylighting” buried streams have the potential to increase N retention. Further work is needed to elucidate the impacts of stream burial on ecosystem functions at the larger stream network scale.     

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.     

Stable N isotope composition of nitrate reflects N transformations during the passage of water through a montane rain forest in Ecuador

Knowledge of the fate of deposited N in the possibly N-limited, highly biodiverse north Andean forests is important because of the possible effects of N inputs on plant performance and species composition. We analyzed concentrations and fluxes of NO₃ ^??CN, NH₄ ⁺?CN and dissolved organic N (DON) in rainfall, throughfall, litter leachate, mineral soil solutions (0.15?C0.30 m depths) and stream water in a montane forest in Ecuador during four consecutive quarters and used the natural ¹⁵N abundance in NO₃ ^? during the passage of rain water through the ecosystem and bulk ??¹⁵N values in soil to detect N transformations. Depletion of ¹⁵N in NO₃ ^? and increased NO₃ ^??CN fluxes during the passage through the canopy and the organic layer indicated nitrification in these compartments. During leaching from the organic layer to mineral soil and stream, NO₃ ^? concentrations progressively decreased and were enriched in ¹⁵N but did not reach the ??¹⁵N values of solid phase organic matter (??¹⁵N = 5.6?C6.7??). This suggested a combination of nitrification and denitrification in mineral soil. In the wettest quarter, the ??¹⁵N value of NO₃ ^? in litter leachate was smaller (??¹⁵N = ?1.58??) than in the other quarters (??¹⁵N = ?9.38 ± SE 0.46??) probably because of reduced mineralization and associated fractionation against ¹⁵N. Nitrogen isotope fractionation of NO₃ ^? between litter leachate and stream water was smaller in the wettest period than in the other periods probably because of a higher rate of denitrification and continuous dilution by isotopically lighter NO₃ ^??CN from throughfall and nitrification in the organic layer during the wettest period. The stable N isotope composition of NO₃ ^? gave valuable indications of N transformations during the passage of water through the forest ecosystem from rainfall to the stream.