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Results for “Journal of Geophysical Research G: Biogeosciences”

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USGS2014

Uncertainty in the fate of soil organic carbon: A comparison of three conceptually different soil decomposition models

Conventional Q10 soil organic matter decomposition models and more complex microbial models are available for making projections of future soil carbon dynamics. However, it is unclear (1) how well the conceptually different approaches can simulate observed decomposition and (2) to what extent the trajectories of long-term simulations differ when using the different approaches. In this study, we compared three structurally different soil carbon (C) decomposition models (one Q10 and two microbial models of different complexity), each with a one- and two-horizon version. The models were calibrated and validated using 4 years of measurements of heterotrophic soil CO 2 efflux from trenched plots in a Dahurian larch ( Larix gmelinii Rupr.) plantation. All models reproduced the observed heterotrophic component of soil CO 2 efflux, but the trajectories of soil carbon dynamics differed substantially in 100 year simulations with and without warming and increased litterfall input, with microbial models that produced better agreement with observed changes in soil organic C in long-term warming experiments. Our results also suggest that both constant and varying carbon use efficiency are plausible when modeling future decomposition dynamics and that the use of a short-term (e.g., a few years) period of measurement is insufficient to adequately constrain model parameters that represent long-term responses of microbial thermal adaption. These results highlight the need to reframe the representation of decomposition models and to constrain parameters with long-term observations and multiple data streams. We urge caution in interpreting future soil carbon responses derived from existing decomposition models because both conceptual and parameter uncertainties are substantial.

Journal of Geophysical Research G: Biogeosciences↗

USGS2015

Source limitation of carbon gas emissions in high-elevation mountain streams and lakes

Inland waters are an important component of the global carbon cycle through transport, storage, and direct emissions of CO 2 and CH 4 to the atmosphere. Despite predictions of high physical gas exchange rates due to turbulent flows and ubiquitous supersaturation of CO 2 —and perhaps also CH 4 —patterns of gas emissions are essentially undocumented for high mountain ecosystems. Much like other headwater networks around the globe, we found that high-elevation streams in Rocky Mountain National Park, USA, were supersaturated with CO 2 during the growing season and were net sources to the atmosphere. CO 2 concentrations in lakes, on the other hand, tended to be less than atmospheric equilibrium during the open water season. CO 2 and CH 4 emissions from the aquatic conduit were relatively small compared to many parts of the globe. Irrespective of the physical template for high gas exchange (high k ), we found evidence of CO 2 source limitation to mountain streams during the growing season, which limits overall CO 2 emissions. Our results suggest a reduced importance of aquatic ecosystems for carbon cycling in high-elevation landscapes having limited soil development and high CO 2 consumption via mineral weathering.

Journal of Geophysical Research G: Biogeosciences↗

USGS2015

Incorporating microbial dormancy dynamics into soil decomposition models to improve quantification of soil carbon dynamics of northern temperate forests

Soil carbon dynamics of terrestrial ecosystems play a significant role in the global carbon cycle. Microbial-based decomposition models have seen much growth recently for quantifying this role, yet dormancy as a common strategy used by microorganisms has not usually been represented and tested in these models against field observations. Here we developed an explicit microbial-enzyme decomposition model and examined model performance with and without representation of microbial dormancy at six temperate forest sites of different forest types. We then extrapolated the model to global temperate forest ecosystems to investigate biogeochemical controls on soil heterotrophic respiration and microbial dormancy dynamics at different temporal-spatial scales. The dormancy model consistently produced better match with field-observed heterotrophic soil CO 2 efflux ( R H ) than the no dormancy model. Our regional modeling results further indicated that models with dormancy were able to produce more realistic magnitude of microbial biomass (<2% of soil organic carbon) and soil R H (7.5 ± 2.4 Pg C yr −1 ). Spatial correlation analysis showed that soil organic carbon content was the dominating factor (correlation coefficient = 0.4–0.6) in the simulated spatial pattern of soil R H with both models. In contrast to strong temporal and local controls of soil temperature and moisture on microbial dormancy, our modeling results showed that soil carbon-to-nitrogen ratio (C:N) was a major regulating factor at regional scales (correlation coefficient = −0.43 to −0.58), indicating scale-dependent biogeochemical controls on microbial dynamics. Our findings suggest that incorporating microbial dormancy could improve the realism of microbial-based decomposition models and enhance the integration of soil experiments and mechanistically based modeling.

Journal of Geophysical Research G: Biogeosciences↗

USGS2016

Wetland tree transpiration modified by river-floodplain connectivity

Hydrologic connectivity provisions water and nutrient subsidies to floodplain wetlands and may be particularly important in floodplains with seasonal water deficits through its effects on soil moisture. In this study, we measured sapflow in 26 trees of two dominant floodplain forest species ( Celtis laevigata and Quercus lyrata ) at two hydrologically distinct sites in the lower White River floodplain in Arkansas, USA. Our objective was to investigate how connectivity-driven water table variations affected water use, an indicator of tree function. Meteorological variables (photosynthetically active radiation and vapor pressure deficit) were the dominant controls over water use at both sites; however, water table variations explained some site differences. At the wetter site, highest sapflow rates were during a late-season overbank flooding event, and no flood stress was apparent. At the drier site, sapflow decreased as the water table receded. The late-season flood pulse that resulted in flooding at the wetter site did not affect the water table at the drier site; accordingly, higher water use was not observed at the drier site. The species generally associated with wetter conditions ( Q. lyrata ) was more positively responsive to the flood pulse. Flood water subsidy lengthened the effective growing season, demonstrating ecological implications of hydrologic connectivity for alleviating water deficits that otherwise reduce function in this humid floodplain wetland.

Arkansas↗

USGS2016

The differing biogeochemical and microbial signatures of glaciers and rock glaciers

Glaciers and rock glaciers supply water and bioavailable nutrients to headwater mountain lakes and streams across all regions of the American West. Here we present a comparative study of the metal, nutrient, and microbial characteristics of glacial and rock glacial influence on headwater ecosystems in three mountain ranges of the contiguous U.S.: The Cascade Mountains, Rocky Mountains, and Sierra Nevada. Several meltwater characteristics (water temperature, conductivity, pH, heavy metals, nutrients, complexity of dissolved organic matter (DOM), and bacterial richness and diversity) differed significantly between glacier and rock glacier meltwaters, while other characteristics (Ca 2+ , Fe 3+ , SiO 2 concentrations, reactive nitrogen, and microbial processing of DOM) showed distinct trends between mountain ranges regardless of meltwater source. Some characteristics were affected both by glacier type and mountain range (e.g. temperature, ammonium (NH 4 + ) and nitrate (NO 3 - ) concentrations, bacterial diversity). Due to the ubiquity of rock glaciers and the accelerating loss of the low latitude glaciers our results point to the important and changing influence that these frozen features place on headwater ecosystems.

California, Colorado, Oregon, Washington, Wyoming↗

USGS2017

Field-scale observations of a transient geobattery resulting from natural attenuation of a crude oil spill

We present evidence of a geobattery associated with microbial degradation of a mature crude oil spill. Self-potential measurements were collected using a vertical array of nonpolarizing electrodes, starting at the land surface and passing through the smear zone where seasonal water table fluctuations have resulted in the coating of hydrocarbons on the aquifer solids. These passive electrical potential measurements exhibit a dipolar pattern associated with a current source. The anodic and cathodic reactions of this natural battery occur below and above the smear zone, respectively. The smear zone is characterized by high magnetic susceptibility values associated with the precipitation of semiconductive magnetic iron phase minerals as a by-product of biodegradation, facilitating electron transfer between the anode and the cathode. This geobattery response appears to have a transient nature, changing on a monthly scale, probably resulting from chemical and physical changes in subsurface conditions such as water table fluctuations.

Journal of Geophysical Research G: Biogeosciences↗

USGS2017

Spatial heterogeneity of within-stream methane concentrations

Streams, rivers, and other freshwater features may be significant sources of CH 4 to the atmosphere. However, high spatial and temporal variabilities hinder our ability to understand the underlying processes of CH 4 production and delivery to streams and also challenge the use of scaling approaches across large areas. We studied a stream having high geomorphic variability to assess the underlying scale of CH 4 spatial variability and to examine whether the physical structure of a stream can explain the variation in surface CH 4 . A combination of high-resolution CH 4 mapping, a survey of groundwater CH 4 concentrations, quantitative analysis of methanogen DNA, and sediment CH 4 production potentials illustrates the spatial and geomorphic controls on CH 4 emissions to the atmosphere. We observed significant spatial clustering with high CH 4 concentrations in organic-rich stream reaches and lake transitions. These sites were also enriched in the methane-producing mcrA gene and had highest CH 4 production rates in the laboratory. In contrast, mineral-rich reaches had significantly lower concentrations and had lesser abundances of mcrA . Strong relationships between CH 4 and the physical structure of this aquatic system, along with high spatial variability, suggest that future investigations will benefit from viewing streams as landscapes, as opposed to ecosystems simply embedded in larger terrestrial mosaics. In light of such high spatial variability, we recommend that future workers evaluate stream networks first by using similar spatial tools in order to build effective sampling programs.

Journal of Geophysical Research G: Biogeosciences↗

USGS2018

Functional group, biomass, and climate change effects on ecological drought in semiarid grasslands

Water relations in plant communities are influenced both by contrasting functional groups (grasses, shrubs) and by climate change via complex effects on interception, uptake and transpiration. We modelled the effects of functional group replacement and biomass increase, both of which can be outcomes of invasion and vegetation management, and climate change on ecological drought (soil water potential below which photosynthesis stops) in 340 semiarid grassland sites over 30‐year periods. Relative to control vegetation (climate and site‐determined mixes of functional groups), the frequency and duration of drought were increased by shrubs and decreased by annual grasses. The rankings of shrubs, control vegetation, and annual grasses in terms of drought effects were generally consistent in current and future climates, suggesting that current differences among functional groups on drought effects predict future differences. Climate change accompanied by experimentally‐increased biomass (i.e. the effects of invasions that increase community biomass, or management that increases productivity through fertilization or respite from grazing) increased drought frequency and duration, and advanced drought onset. Our results suggest that the replacement of perennial temperate semiarid grasslands by shrubs, or increased biomass, can increase ecological drought both in current and future climates.

Journal of Geophysical Research G: Biogeosciences↗

USGS2018

On factors influencing air-water gas exchange in emergent wetlands

Knowledge of gas exchange in wetlands is important in order to determine fluxes of climatically and biogeochemically important trace gases and to conduct mass balances for metabolism studies. Very few studies have been conducted to quantify gas transfer velocities in wetlands, and many wind speed/gas exchange parameterizations used in oceanographic or limnological settings are inappropriate under conditions found in wetlands. Here six measurements of gas transfer velocities are made with SF 6 tracer release experiments in three different years in the Everglades, a subtropical peatland with surface water flowing through emergent vegetation. The experiments were conducted under different flow conditions and with different amounts of emergent vegetation to determine the influence of wind, rain, water flow, waterside thermal convection, and vegetation on air-water gas exchange in wetlands. Measured gas transfer velocities under the different conditions ranged from 1.1 cm h −1 during baseline conditions to 3.2 cm h −1 when rain and water flow rates were high. Commonly used wind speed/gas exchange relationships would overestimate the gas transfer velocity by a factor of 1.2 to 6.8. Gas exchange due to thermal convection was relatively constant and accounted for 14 to 51% of the total measured gas exchange. Differences in rain and water flow among the different years were responsible for the variability in gas exchange, with flow accounting for 37 to 77% of the gas exchange, and rain responsible for up to 40%.

Florida↗

USGS2013

Nitrate in watersheds: straight from soils to streams?

Human activities are rapidly increasing the global supply of reactive N and substantially altering the structure and hydrologic connectivity of managed ecosystems. There is long-standing recognition that N must be removed along hydrologic flowpaths from uplands to streams, yet it has proven difficult to assess the generality of this removal across ecosystem types, and whether these patterns are influenced by land-use change. To assess how well upland nitrate (NO 3 - ) loss is reflected in stream export, we gathered information from >50 watershed biogeochemical studies that reported nitrate concentrations ([NO 3 - ]) for stream water and for either upslope soil solution or groundwater NO 3 - to examine whether stream export of NO3- accurately reflects upland NO 3 - losses. In this dataset, soil solution and streamwater [NO 3 - ] were correlated across 40 undisturbed forest watersheds, with streamwater [NO 3 - ] typically half (median = 50%) soil solution [NO 3 - ]. A similar relationship was seen in 10 disturbed forest watersheds. However, for 12 watersheds with significant agricultural or urban development, the intercept and slope were both significantly higher than the relationship seen in forest watersheds. Differences in concentration between soil solution or groundwater and stream water may be attributed to biological uptake, microbial processes including denitrification, and/or preferential flow routing. The results of this synthesis are consistent with the hypotheses that undisturbed watersheds have a significant capacity to remove nitrate after it passes below the rooting zone and that land use changes tend to alter the efficiency or the length of watershed flowpaths, leading to reductions in nitrate removal and increased stream nitrate concentrations.

Journal of Geophysical Research G: Biogeosciences↗

USGS2013

Emissions of carbon dioxide and methane from a headwater stream network of interior Alaska

Boreal ecosystems store significant quantities of organic carbon (C) that may be vulnerable to degradation as a result of a warming climate. Despite their limited coverage on the landscape, streams play a significant role in the processing, gaseous emission, and downstream export of C, and small streams are thought to be particularly important because of their close connection with the surrounding landscape. However, ecosystem carbon studies do not commonly incorporate the role of the aquatic conduit. We measured carbon dioxide (CO 2 ) and methane (CH 4 ) concentrations and emissions in a headwater stream network of interior Alaska underlain by permafrost to assess the potential role of stream gas emissions in the regional carbon balance. First-order streams exhibited the greatest variability in fluxes of CO 2 and CH 4, and the greatest mean p CO 2 . High-resolution time series of stream p CO 2 and discharge at two locations on one first-order stream showed opposing p CO 2 responses to storm events, indicating the importance of hydrologic flowpaths connecting CO 2 -rich soils with surface waters. Repeated longitudinal surveys on the stream showed consistent areas of elevated p CO 2 and p CH 4 , indicative of discrete hydrologic flowpaths delivering soil water and groundwater having varying chemistry. Up-scaled basin estimates of stream gas emissions suggest that streams may contribute significantly to catchment-wide CH 4 emissions. Overall, our results indicate that while stream-specific gas emission rates are disproportionately high relative to the terrestrial landscape, both stream surface area and catchment normalized emission rates were lower than those documented for the Yukon River Basin as a whole. This may be due to limitations of C sources and/or C transport to surface waters.

Journal of Geophysical Research G: Biogeosciences↗

USGS2013

Hydrologic controls on the transport and cycling of carbon and nitrogen in a boreal catchment underlain by continuous permafrost

Boreal ecosystems represent a large carbon (C) reservoir and a substantial source of greenhouse gases. Hydrologic conditions dictate whether C leached from boreal soils is processed in catchments or flushed to less productive environments via the stream. This study quantified hydrologic and biogeochemical C loss from a boreal catchment underlain by frozen silt, where flowpaths may deepen as the active layer thaws over the summer. We hypothesized a decrease in the magnitude of C mineralization over the summer associated with changing flowpaths and decreasing hydrologic connectivity, organic matter lability, and nitrogen (N) availability. Conservative tracers were used to partition C and N loss between catchment export and biogeochemical processing. Coupling tracers with tributary and porewater chemistry indicated C and N cycling in soil flowpaths, with an exponential decrease over the summer. Nitrate was primarily reduced in hillslope flowpaths and the lack of N reaching the stream appeared to limit C mineralization. Stream export accounted for the greatest loss of C, removing 247 and 113 mol hr −1 in the early and late summer, respectively. Reactivity was related to hydrologic connectivity between the soils and stream, which was greatest early in the summer and following a large flood. While a warming climate may increase storage potential in thawed soils, the early‐season flush of labile material and late‐season runoff through mineral flowpaths may maintain high C export rates. Therefore, we highlight physical export as a dominant cause of aqueous C loss from silty catchments as the Arctic continues to thaw.

Journal of Geophysical Research G: Biogeosciences↗

USGS2006

Effect of tree line advance on carbon storage in NW Alaska

We investigated the size, distribution, and temporal dynamics of ecosystem carbon (C) pools in an area of recent tree line advance, northwest Alaska. Repeat aerial photographs show forest cover increased ???10% in our study area since 1949. We sampled C pools of four principal ecosystem types, tussock tundra, shrub tundra, woodland, and forest, all located on a 600-800 year old river terrace. Significant differences between ecosystem C pools, both above ground and below ground existed. Tundra sites store >22.2 kg C/m2, shrub tundra sites and woodland sites store 9.7 kg C/m2 and 14.3 kg C/m2, respectively, and forest sites store 14.4 kg C/m2. Landscape variation of total ecosystem C was primarily due to organic soil C and was secondarily due to C stored in trees. Soil C/N profiles of shrub tundra sites and woodland sites showed similarities with forest site soils at surface and tundra site soils at depth. We hypothesize that tundra systems transformed to forest systems in this area under a progression of permafrost degradation and enhanced drainage. On the basis of C pool estimates for the different ecosystem types, conversion of tundra sites to forest may have resulted in a net loss of > 7.8 kg C/m2, since aboveground C gains were more than offset by belowground C losses to decomposition in the tundra sites. Tree line advance therefore might not increase C storage in high-latitude ecosystems and thus might not, as previously suggested, act as a negative feedback to warming. Key to this hypothesis and to its projection to future climate response is the fate of soil carbon upon warming and permafrost drainage. Copyright 2006 by the American Geophysical Union.

Journal of Geophysical Research G: Biogeosciences↗

USGS2006

Geomorphic control of landscape carbon accumulation

We use the CREEP process-response model to simulate soil organic carbon accumulation in an undisturbed prairie site in Iowa. Our primary objectives are to identify spatial patterns of carbon accumulation, and explore the effect of erosion on basin-scale C accumulation. Our results point to two general findings. First, redistribution of soil carbon by erosion results in a net increase in basin-wide carbon storage relative to a noneroding environment. Landscape-average mean residence times are increased in an eroding landscape owing to the burial/preservation of otherwise labile C. Second, field observations taken along a slope transect may overlook significant intraslope variations in carbon accumulation. Spatial patterns of modeled deep C accumulation are complex. While surface carbon with its relatively short equilibration time is predictable from surface properties, deep carbon is strongly influenced by the landscape's geomorphic and climatic history, resulting in wide spatial variability. Convergence and divergence associated with upland swales and interfluves result in bimodal carbon distributions in upper and mid slopes; variability in carbon storage within modeled mid slopes was as high as simulated differences between erosional shoulders and depositional valley bottoms. The bimodality of mid-slope C variability in the model suggests that a three-dimensional sampling strategy is preferable over the traditional two-dimensional analog or "catena" approach. Copyright 2006 by the American Geophysical Union.

Journal of Geophysical Research G: Biogeosciences↗

USGS2006

Modeling physical and biogeochemical controls over carbon accumulation in a boreal forest soil

Boreal soils are important to the global C cycle owing to large C stocks, repeated disturbance from fire, and the potential for permafrost thaw to expose previously stable, buried C. To evaluate the primary mechanisms responsible for both short- and long-term C accumulation in boreal soils, we developed a multi-isotope (12,14C) Soil C model with dynamic soil layers that develop through time as soil organic matter burns and reaccumulates. We then evaluated the mechanisms that control organic matter turnover in boreal regions including carbon input rates, substrate recalcitrance, soil moisture and temperature, and the presence of historical permafrost to assess the importance of these factors in boreal C accumulation. Results indicate that total C accumulation is controlled by the rate of carbon input, decomposition rates, and the presence of historical permafrost. However, unlike more temperate ecosystems, one of the key mechanisms involved in C preservation in boreal soils examined here is the cooling of subsurface soil layers as soil depth increases rather than increasing recalcitrance in subsurface soils. The propagation of the 14C bomb spike into soils also illustrates the importance of historical permafrost and twentieth century warming in contemporary boreal soil respiration fluxes. Both 14C and total C simulation data also strongly suggest that boreal SOM need not be recalcitrant to accumulate; the strong role of soil temperature controls on boreal C accumulation at our modeling test site in Manitoba, Canada, indicates that carbon in the deep organic soil horizons is probably relatively labile and thus subject to perturbations that result from changing climatic conditions in the future. Copyright 2006 by the American Geophysical Union.

Journal of Geophysical Research G: Biogeosciences↗

USGS2006

Effects of permafrost melting on CO2 and CH4 exchange of a poorly drained black spruce lowland

[1] Permafrost melting is occurring in areas of the boreal forest region where large amounts of carbon (C) are stored in organic soils. We measured soil respiration, net CO 2 flux, and net CH 4 flux during May–September 2003 and March 2004 in a black spruce lowland in interior Alaska to better understand how permafrost thaw in poorly drained landscapes affects land‐atmosphere CO 2 and CH 4 exchange. Sites included peat soils underlain by permafrost at ∼0.4 m depth (permafrost plateau, PP), four thermokarst wetlands (TW) having no permafrost in the upper 2.2 m, and peat soils bordering the thermokarst wetlands having permafrost at ∼0.5 m depth (thermokarst edges, TE). Soil respiration rates were not significantly different among the sites, and 5‐cm soil temperature explained 50–91% of the seasonal variability in soil respiration within the sites. Groundcover vegetation photosynthesis (calculated as net CO 2 minus soil respiration) was significantly different among the sites (TW > TE > PP), which can be partly attributed to the difference in photosynthetically active radiation reaching the ground at each site type. Methane emission rates were 15 to 28 times greater from TW than from TE and PP. We modeled annual soil respiration and groundcover vegetation photosynthesis using soil temperature and radiation data, and CH 4 flux by linear interpolation. We estimated all sites as net C gas sources to the atmosphere (not including tree CO 2 uptake at PP and TE), although the ranges in estimates when accounting for errors were large enough that TE and TW may have been net C sinks.

Journal of Geophysical Research G: Biogeosciences↗

USGS2006

Shrinking ponds in subarctic Alaska based on 1950-2002 remotely sensed images

Over the past 50 years, Alaska has experienced a warming climate with longer growing seasons, increased potential evapotranspiration, and permafrost warming. Research from the Seward Peninsula and Kenai Peninsula has demonstrated a substantial landscape-level trend in the reduction of surface water and number of closed-basin ponds. We investigated whether this drying trend occurred at nine other regions throughout Alaska. One study region was from the Arctic Coastal Plain where depp permafrost occurs continuously across the landscape. The other eight study regions were from the boreal forest regions where discontinuous permafrost occurs. Mean annual precipitation across the study regions ranged from 100 to over 700 min yr-1. We used remotely sensed imagery from the 1950s to 2002 to inventory over 10,000 closed-basin ponds from at least three periods from this time span. We found a reduction in the area and number of shallow, closed-basin ponds for all boreal regions. In contrast, the Arctic Coastal Plain region had negligible change in the area of closed-basin ponds. Since the 1950s, surface water area of closed-basin ponds included in this analysis decreased by 31 to 4 percent, and the total number of closed-basin ponds surveyed within each study region decreased from 54 to 5 percent. There was a significant increasing trend in annual mean temperature and potential evapotranspiration since the 1950s for all study regions. There was no significant trend in annual precipitation during the same period. The regional trend of shrinking ponds may be due to increased drainage as permafrost warms, or increased evapotranspiration during a warmer and extended growing season. Copyright 2006 by the American Geophysical Union.

Journal of Geophysical Research G: Biogeosciences↗

USGS2006

Quantity-activity relationship of denitrifying bacteria and environmental scaling in streams of a forested watershed

The spatial variability of subreach denitrification rates in streams was evaluated with respect to controlling environmental conditions, molecular examination of denitrifying bacteria, and dimensional analysis. Denitrification activities ranged from 0 and 800 ng-N gsed-1 d-1 with large variations observed within short distances (<50 m) along stream reaches. A log-normal probability distribution described the range in denitrification activities and was used to define low (16% of the probability distributibn), medium (68%), and high (16%) denitrification potential groups. Denitrifying bacteria were quantified using a competitive polymerase chain reaction (cPCR) technique that amplified the nirK gene that encodes for nitrite reductase. Results showed a range of nirK quantities from 103 to 107 gene-copy-number gsed.-1 A nonparametric statistical test showed no significant difference in nirK quantifies among stream reaches, but revealed that samples with a high denitrification potential had significantly higher nirK quantities. Denitrification activity was positively correlated with nirK quantities with scatter in the data that can be attributed to varying environmental conditions along stream reaches. Dimensional analysis was used to evaluate denitrification activities according to environmental variables that describe fluid-flow properties, nitrate and organic material quantities, and dissolved oxygen flux. Buckingham's pi theorem was used to generate dimensionless groupings and field data were used to determine scaling parameters. The resulting expressions between dimensionless NO3- flux and dimensionless groupings of environmental variables showed consistent scaling, which indicates that the subreach variability in denitrification rates can be predicted by the controlling physical, chemical, and microbiological conditions. Copyright 2006 by the American Geophysical Union.

Journal of Geophysical Research G: Biogeosciences↗