Nitrogen critical loads for alpine vegetation and terrestrial ecosystem response: are we there yet?

Increases in the deposition of anthropogenic nitrogen (N) have been linked to several terrestrial ecological changes, including soil biogeochemistry, plant stress susceptibility, and community diversity. Recognizing the need to identify sensitive indicators of biotic response to N deposition, we empirically estimated the N critical load for changes in alpine plant community composition and compared this with the estimated critical load for soil indicators of ecological change. We also measured the degree to which alpine vegetation may serve as a sink for anthropogenic N and how much plant sequestration is related to changes in species composition. We addressed these research goals by adding 20, 40, or 60 kg N x ha(-1) x yr(-1), along with an ambient control (6 kg N x ha(-1) x yr(-1) total deposition), to a species-rich alpine dry meadow for an eight-year period. Change in plant species composition associated with the treatments occurred within three years of the initiation of the experiment and were significant at all levels of N addition. Using individual species abundance changes and ordination scores, we estimated the N critical loads (total deposition) for (1) change in individual species to be 4 kg N x ha(-1) yr(-1) and (2) for overall community change to be 10 kg N x ha(-1) x yr(-1). In contrast, increases in NO3- leaching, soil solution inorganic NO3-, and net N nitrification occurred at levels above 20 kg N x ha(-1) x yr(-1). Increases in total aboveground biomass were modest and transient, occurring in only one of the three years measured. Vegetative uptake of N increased significantly, primarily as a result of increasing tissue N concentrations and biomass increases in subdominant species. Aboveground vegetative uptake of N accounted for <40% of the N added. The results of this experiment indicate that changes in vegetation composition will precede detectable changes in more traditionally used soil indicators of ecosystem responses to N deposition and that changes in species composition are probably ongoing in alpine dry meadows of the Front Range of the Colorado Rocky Mountains. Feedbacks to soil N cycling associated with changes in litter quality and species composition may result in only short-term increases in vegetation N pools.     

Greenhouse gas fluxes in southeastern U.S. coastal plain wetlands under contrasting land uses

Whether through sea level rise or wetland restoration, agricultural soils in coastal areas will be inundated at increasing rates, renewing connections to sensitive surface waters and raising critical questions about environmental trade-offs. Wetland restoration is often implemented in agricultural catchments to improve water quality through nutrient removal. Yet flooding of soils can also increase production of the greenhouse gases nitrous oxide and methane, representing a potential environmental trade-off. Our study aimed to quantify and compare greenhouse gas emissions from unmanaged and restored forested wetlands, as well as actively managed agricultural fields within the North Carolina coastal plain, USA. In sampling conducted once every two months over a two-year comparative study, we found that soil carbon dioxide flux (range: 8000-64 800 kg CO2 x ha(-1) x yr(-1)) comprised 66-100% of total greenhouse gas emissions from all sites and that methane emissions (range: -6.87 to 197 kg CH4 x ha(-1) x yr(-1)) were highest from permanently inundated sites, while nitrous oxide fluxes (range: -1.07 to 139 kg N2O x ha(-1) x yr(-1)) were highest in sites with lower water tables. Contrary to predictions, greenhouse gas fluxes (as CO2 equivalents) from the restored wetland were lower than from either agricultural fields or unmanaged forested wetlands. In these acidic coastal freshwater ecosystems, the conversion of agricultural fields to flooded young forested wetlands did not result in increases in greenhouse gas emissions.     

Land use change and nitrogen enrichment of a Rocky Mountain watershed.

Headwater ecosystems may have a limited threshold for retaining and removing nutrients delivered by certain types of land use. Nitrogen enrichment was studied in a Rocky Mountain watershed undergoing rapid expansion of population and residential development. Study sites were located along a 30-km transect from the headwaters of the Blue River to Lake Dillon, a major source of drinking water for Denver, Colorado. Ground water in residential areas with septic systems showed high concentrations of nitrate-N (4.96 +/- 1.22 mg/L, mean +/- SE), and approximately 40% of wells contained nitrate with delta15N values in the range of wastewater. Concentrations of dissolved inorganic nitrogen (DIN) in tributaries with residential development peaked during spring snowmelt as concentrations of DIN declined to below detection limits in undeveloped tributaries. Annual export of dissolved organic nitrogen (DON) was considerably lower in residential streams, suggesting a change in forms of N with development. The seasonal delta15N of algae in residential streams was intermediate between baseline values from undeveloped streams and stream algae grown on wastewater. Between 19% and 23% of the annual N export from developed tributaries was derived from septic systems, as estimated from the delta15N of algae. This range was similar to the amount of N export above background determined independently from mass-balance estimates. From a watershed perspective, total loading of N to the Blue River catchment from septic and municipal wastewater (2 kg x ha(-1) x yr(-1)) is currently less than the amount from background atmospheric sources (3 kg x ha(-1) x yr(-1)). Nonetheless, nitrate-N concentrations exceeded limits for safe drinking water in some groundwater wells (10 mg/L), residential streams showed elevated seasonal patterns of nitrate-N concentration and ratios of DIN to total dissolved phosphorus, and seasonal minimum concentrations of nitrate-N in Lake Dillon have increased exponentially to 80 microg/L over the last decade from an initial value near zero. Results suggest that isotopic ratios in autotrophs can be used to detect and quantify increases in N enrichment associated with land use change. The biotic capacity of headwater ecosystems to assimilate increases in inorganic N from residential development may be insufficient to prevent nitrogen enrichment over considerable distances and multiple aquatic ecosystems downstream.     

Empirical modeling of atmospheric deposition in mountainous landscapes.

Atmospheric deposition has long been recognized as an important source of pollutants and nutrients to ecosystems. The need for reliable, spatially explicit estimates of total atmospheric deposition (wet + dry + cloud) is central, not only to air pollution effects researchers, but also for calculation of input-output budgets, and to decision makers faced with the challenge of assessing the efficacy of policy initiatives related to deposition. Although atmospheric deposition continues to represent a critical environmental and scientific issue, current estimates of total deposition have large uncertainties, particularly across heterogeneous landscapes such as montane regions. We developed an empirical modeling approach that predicts total deposition as a function of landscape features. We measured indices of total deposition to the landscapes of Acadia (121 km2) and Great Smoky Mountains (2074 km2) National Parks (USA). Using approximately 300-400 point measurements and corresponding landscape variables at each park, we constructed a statistical (general linear) model relating the deposition index to landscape variables measured in the field. The deposition indices ranged over an order of magnitude, and in response to vegetation type and elevation, which together explained approximately 40% of the variation in deposition. Then, using the independent landscape variables available in GIS data layers, we created a GIS-relevant statistical nitrogen (N) and sulfur (S) deposition model (LandMod). We applied this model to create park-wide maps of total deposition that were scaled to wet and dry deposition data from the closest national network monitoring stations. The resultant deposition maps showed high spatial heterogeneity and a four- to sixfold variation in "hot spots" and "cold spots" of N and S deposition ranging from 3 to 31 kg N x ha(-1) x yr(-1) and from 5 to 42 kg S x ha(-1) x yr(-1) across these park landscapes. Area-weighted deposition was found to be up to 70% greater than NADP plus CASTNET monitoring-station estimates together. Model-validation results suggest that the model slightly overestimates deposition for deciduous and coniferous forests at low elevation and underestimates deposition for high-elevation coniferous forests. The spatially explicit deposition estimates derived from LandMod are an improvement over what is currently available. Future research should test LandMod in other mountainous environments and refine it to account for (currently) unexplained variation in deposition.     

Carbon sequestration in California agriculture, 1980-2000.

To better understand agricultural carbon fluxes in California, USA, we estimated changes in soil carbon and woody material between 1980 and 2000 on 3.6 x 10(6) ha of farmland in California. Combining the CASA (Carnegie-Ames-Stanford Approach) model with data on harvest indices and yields, we calculated net primary production, woody production in orchard and vineyard crops, and soil carbon. Over the 21-yr period, two trends resulted in carbon sequestration. Yields increased an average of 20%, corresponding to greater plant biomass and more carbon returned to the soils. Also, orchards and vineyards increased in area from 0.7 x 10(6) ha to 1.0 x 10(6) ha, displacing field crops and sequestering woody carbon. Our model estimates that California's agriculture sequestered an average of 19 g C x m(-2) x yr(-1). Sequestration was lowest in non-rice annual cropland, which sequestered 9 g C x m(-2) x yr(-1) of soil carbon, and highest on land that switched from annual cropland to perennial cropland. Land that switched from annual crops to vineyards sequestered 68 g C x m(-2) x yr(-1), and land that switched from annual crops to orchards sequestered 85 g C x m(-2) x yr(-1). Rice fields, because of a reduction in field burning, sequestered 55 g C x m(-2) x yr(-1) in the 1990s. Over the 21 years, California's 3.6 x 10(6) ha of agricultural land sequestered 11.0 Tg C within soils and 3.5 Tg C in woody biomass, for a total of 14.5 Tg C statewide. This is equal to 0.7% of the state's total fossil fuel emissions over the same time period. If California's agriculture adopted conservation tillage, changed management of almond and walnut prunings, and used all of its orchard and vineyard waste wood in the biomass power plants in the state, California's agriculture could offset up to 1.6% of the fossil fuel emissions in the state.     

Birds transport nutrients to fragmented forests in an urban landscape.

The influence of urbanization on nutrient cycling is vaguely known. Here we document that birds, especially those increasing in urban areas (such as crows, Corvus macrorhynchos and C. corone), affect nutrient cycles. Using fecal traps, we measured phosphorus (P) and nitrogen (N) input from the excrement of birds in fragmented forests in an urban landscape. Sources of avian feces were examined on the basis of carbon (C), N, and P percentages and stable isotopes of delta15N and delta13C. Nitrogen and P input was aggregated in the urban landscape, being especially high at the forest where crows roosted during winter. The annual P input due to bird droppings (range 0.068-0.460 kg x ha(-1) x yr(-1); mean 0.167 kg x ha(-1) x yr(-1)) was 12.4% of the total of other pathways in typical forests and 52.9% in the evergreen forest where crows roosted. The annual N input due to bird droppings (range 0.44-3.49 kg x ha(-1) x yr(-1); mean 1.15 kg x ha(-1) x yr(-1)) was 5.2% of the total of other pathways in typical forests and 27.0% in the evergreen forest used by roosting crows. Expected sources of nutrients in feces included insects in the breeding season, fruits in autumn, and mammals and birds in winter. Stable isotopes suggested that the source of nutrients in forests used by roosting crows was from outside the forest. Therefore, birds played a significant role as transporters of nutrients from garbage (including fish, livestock, and/or C4 plants such as corn, with high delta15N and delta13C) in residential and business areas to fragmented evergreen forests, especially near their winter roosts.     

Global change shifts vegetation and plant-parasite interactions in a boreal mire

The aim of this study was to detect vegetation change and to examine trophic interactions in a Sphagnum-dominated mire in response to raised temperature and nitrogen (N) addition. A long-term global-change experiment was established in 1995, with monthly additions of N (30 kg x ha(-1) x yr(-1)) and sulfur (20 kg x ha(-1) x yr(-1)) during the vegetation period. Mean air temperature was raised by 3.6 degrees C with warming chambers. Vegetation responses were negligible for all treatments for the first four years, and no sulfur effect was seen during the course of the experiment. However, after eight years of continuous treatments, the closed Sphagnum carpet was drastically reduced from 100% in 1995 down to 41%, averaged over all N-treated plots. Over the same period, total vascular plant cover (of the graminoid Eriophorum vaginatum and the two dwarf-shrubs Andromeda polifolia and Vaccinium oxycoccos) increased from 24% to an average of 70% in the N plots. Nitrogen addition caused leaf N concentrations to rise in the two dwarf-shrubs, while for E. vaginatum, leaf N remained unchanged, indicating that the graminoid to a larger extent than the dwarf-shrubs allocated supplemented N to growth. Concurrent with foliar N accumulation of the two dwarf-shrubs, we observed increased disease incidences caused by parasitic fungi, with three species out of 16 showing a significant increase. Warming caused a significant decrease in occurrence of three parasitic fungal species. In general, decreased disease incidences were found in temperature treatments for A. polifolia and in plots without N addition for V. oxycoccos. The study demonstrates that both bryophytes and vascular plants at boreal mires, only receiving background levels of nitrogen of about 2 kg x ha(-1) x yr(-1), exhibit a time lag of more than five years in response to nitrogen and temperature rise, emphasizing the need for long-term experiments. Moreover, it shows that trophic interactions are likely to differ markedly in response to climate change and increased N deposition, and that these interactions might play an important role in controlling the change in mire vegetation composition, with implications for both carbon sequestration and methane emission.     

Sinks for nitrogen inputs in terrestrial ecosystems: a meta-analysis of 15N tracer field studies

Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (< 1 week after 15N tracer application), total ecosystem 15N recovery was negatively correlated with fine-root and soil 15N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3-18 months after 15N tracer application), total ecosystem 15N retention was negatively correlated with foliar natural-abundance 15N but was positively correlated with mineral soil C and N concentration and C:N, showing that plant and soil natural-abundance 15N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem 15N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for 15N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N x ha(-1) x yr(-1) above which most ecosystems showed net losses of applied 15N tracer in response to N fertilizer addition.     

Biogeochemistry of a temperate forest nitrogen gradient

Wide natural gradients of soil nitrogen (N) can be used to examine fundamental relationships between plant-soil-microbial N cycling and hydrologic N loss, and to test N-saturation theory as a general framework for understanding ecosystem N dynamics. We characterized plant production, N uptake and return in litterfall, soil gross and net N mineralization rates, and hydrologic N losses of nine Douglas-fir (Pseudotsuga menziesii) forests across a wide soil N gradient in the Oregon Coast Range (U.S.A.). Surface mineral soil N (0-10 cm) ranged nearly three-fold from 0.29% to 0.78% N, and in contrast to predictions of N-saturation theory, was linearly related to 10-fold variation in net N mineralization, from 8 to 82 kg N.ha(-1) x yr(-1). Net N mineralization was unrelated to soil C:N, soil texture, precipitation, and temperature differences among sites. Net nitrification was negatively related to soil pH, and accounted for <20% of net N mineralization at low-N sites, increasing to 85-100% of net N mineralization at intermediate- and high-N sites. The ratio of net: gross N mineralization and nitrification increased along the gradient, indicating progressive saturation of microbial N demands at high soil N. Aboveground N uptake by plants increased asymptotically with net N mineralization to a peak of approximately 35 kg N.ha(-1) x yr(-1). Aboveground net primary production per unit net N mineralization varied inversely with soil N, suggesting progressive saturation of plant N demands at high soil N. Hydrologic N losses were dominated by dissolved organic N at low-N sites, with increased nitrate loss causing a shift to dominance by nitrate at high-N sites, particularly where net nitrification exceeded plant N demands. With the exception of N mineralization patterns, our results broadly support the application of the N-saturation model developed from studies of anthropogenic N deposition to understand N cycling and saturation of plant and microbial sinks along natural soil N gradients. This convergence of behavior in unpolluted and polluted forest N cycles suggests that where future reductions in deposition to polluted sites do occur, symptoms of N saturation are most likely to persist where soil N content remains elevated.     

Plant diversity, CO2, and N influence inorganic and organic N leaching in grasslands

In nitrogen (N)-limited systems, the potential to sequester carbon depends on the balance between N inputs and losses as well as on how efficiently N is used, yet little is known about responses of these processes to changes in plant species richness, atmospheric CO2 concentration ([CO2]), and N deposition. We examined how plant species richness (1 or 16 species), elevated [CO2] (ambient or 560 ppm), and inorganic N addition (0 or 4 g x m(-2) x yr(-1)) affected ecosystem N losses, specifically leaching of dissolved inorganic N (DIN) and organic N (DON) in a grassland field experiment in Minnesota, USA. We observed greater DIN leaching below 60 cm soil depth in the monoculture plots (on average 1.8 and 3.1 g N x m(-2) x yr(-1) for ambient N and N-fertilized plots respectively) than in the 16-species plots (0.2 g N x m(-2) x yr(-1) for both ambient N and N-fertilized plots), particularly when inorganic N was added. Most likely, loss of complementary resource use and reduced biological N demand in the monoculture plots caused the increase in DIN leaching relative to the high-diversity plots. Elevated [CO2] reduced DIN concentrations under conditions when DIN concentrations were high (i.e., in N-fertilized and monoculture plots). Contrary to the results for DIN, DON leaching was greater in the 16-species plots than in the monoculture plots (on average 0.4 g N x m(-2) x yr(-1) in 16-species plots and 0.2 g N x m(-2) x yr(-1) in monoculture plots). In fact, DON dominated N leaching in the 16-species plots (64% of total N leaching as DON), suggesting that, even with high biological demand for N, substantial amounts of N can be lost as DON. We found no significant main effects of elevated [CO2] on DIN or DON leaching; however, elevated [CO2] reduced the positive effect of inorganic N addition on DON leaching, especially during the second year of observation. Our results suggest that plant species richness, elevated [CO2], and N deposition alter DIN loss primarily through changes in biological N demand. DON losses can be as large as DIN loss but are more sensitive to organic matter production and turnover.     

Denitrification mitigates N flux through the stream-floodplain complex of a desert city

The Indian Bend Wash (IBW) flood-control project relies on a greenbelt to carry floods through Scottsdale, Arizona, USA. The greenbelt is characterized by a chain of shallow artificial lakes in a larger floodplain of irrigated turf, which has been protected from encroaching urban development. As such, this urban stream-floodplain complex can be divided into three subsystems: artificial lakes, channelized stream segments, and floodplain. We conducted experiments to evaluate which, if any, of these subsystems were important sites of denitrification, and to explore factors controlling denitrification rates. Denitrification enzyme activity (DEA) bioassays were conducted on sediments from eight lake and six stream segments as well as soil samples from eight floodplain transects. Mass-specific potential denitrification rates were significantly higher in lakes than in streams or floodplains. Nutrient limitation bioassays revealed that nitrate (NO3-) limited denitrification in lake sediments, a surprising finding given that NO3(-)-rich groundwater additions frequently raised lake NO3(-) concentration above 1 mg N/L. Experiments on intact lake cores suggested that denitrification was limited by the rate NO3(-) diffused into sediments, rather than its availability in overlying water. Floodplain denitrification was limited by water content, not NO3(-) or C, and irrigation of soils stimulated denitrification. We constructed a N budget for the IBW stream-floodplain complex based on our experimental results. We found that both lakes and floodplains removed large quantities of N, with denitrification removing 261 and 133 kg N ha(-1) yr(-1) from lake sediments and floodplain soils, respectively, indicating that lakes are hotspots for denitrification. Nevertheless, because floodplain area was >4.5 times that of lakes, floodplain soils removed nearly 2.5 times as much N as lake sediments. Given the desert's low annual precipitation, a finding that floodplain soils are active sites of denitrification might seem implausible; however, irrigation is common in urban landscapes, and it elevated annual denitrification in IBW. Based on our results, we conclude that construction of artificial lakes created hotspots while application of irrigation water created hot moments for denitrification in the stream-floodplain complex, demonstrating that management decisions can improve the ability of urban streams to provide critical ecosystem services like N retention.     

Atmospheric deposition, mineralization and leaching of nitrogen in subtropical forested catchments, South China

In recent years, China has conducted considerable research focusing on the emission and effects of sulphur (S) on human health and ecosystems. By contrast, there has been little emphasis on anthropogenic nitrogen (N) so far, even though studies conducted abroad indicate that long-range atmospheric transport of N and ecological effects (e.g. acidification of soil and water) may be significant. The Sino-Norwegian project IMPACTS, launched in 1999, has established monitoring sites at five forest ecosystems in the southern part of PR China to collect comprehensive data on air quality, acidification status and ecological effects. Here we present initial results about N dynamics at two of the IMPACTS sites located near Chongqing and Changsha, including estimation of atmospheric deposition fluxes of NOx and NHx and soil N transformations. Nitrogen deposition is high at both sites when compared with values from Europe and North America (25-38 kg ha(-1) yr(-1)). About 70% of the deposited N comes as NH4, probably derived from agriculture. Leaching of N from soils is high and nearly all as NO3-. Transformation of N to NO3- in soils results in acidification rates that are high compared to rates found elsewhere. Despite considerable leaching of NO3- from the root zone of the soils, little NO3- appears in streamwater. This indicates that N retention or denitrification, both causing acid neutralization, may be important and probably occur in the groundwater and groundwater discharge zones. The soil flux density of mineral N, which is the sum of N deposition and N mineralization, and which is dominated by the N mineralization flux, may be a good indicator for leaching of NO3- in soils. However, this indicator seems site specific probably due to differences in land-use history and current N requirement.     

Life-cycle assessment of net greenhouse-gas flux for bioenergy cropping systems.

Bioenergy cropping systems could help offset greenhouse gas emissions, but quantifying that offset is complex. Bioenergy crops offset carbon dioxide emissions by converting atmospheric CO2 to organic C in crop biomass and soil, but they also emit nitrous oxide and vary in their effects on soil oxidation of methane. Growing the crops requires energy (e.g., to operate farm machinery, produce inputs such as fertilizer) and so does converting the harvested product to usable fuels (feedstock conversion efficiency). The objective of this study was to quantify all these factors to determine the net effect of several bioenergy cropping systems on greenhouse-gas (GHG) emissions. We used the DAYCENT biogeochemistry model to assess soil GHG fluxes and biomass yields for corn, soybean, alfalfa, hybrid poplar, reed canarygrass, and switchgrass as bioenergy crops in Pennsylvania, USA. DAYCENT results were combined with estimates of fossil fuels used to provide farm inputs and operate agricultural machinery and fossil-fuel offsets from biomass yields to calculate net GHG fluxes for each cropping system considered. Displaced fossil fuel was the largest GHG sink, followed by soil carbon sequestration. N20 emissions were the largest GHG source. All cropping systems considered provided net GHG sinks, even when soil C was assumed to reach a new steady state and C sequestration in soil was not counted. Hybrid poplar and switchgrass provided the largest net GHG sinks, >200 g CO2e-C x m(-2) x yr(-1) for biomass conversion to ethanol, and >400 g CO2e-C x m(-2) x yr(-1) for biomass gasification for electricity generation. Compared with the life cycle of gasoline and diesel, ethanol and biodiesel from corn rotations reduced GHG emissions by approximately 40%, reed canarygrass by approximately 85%, and switchgrass and hybrid poplar by approximately 115%.     

Denitrification across landscapes and waterscapes: a synthesis.

Denitrification is a critical process regulating the removal of bioavailable nitrogen (N) from natural and human-altered systems. While it has been extensively studied in terrestrial, freshwater, and marine systems, there has been limited communication among denitrification scientists working in these individual systems. Here, we compare rates of denitrification and controlling factors across a range of ecosystem types. We suggest that terrestrial, freshwater, and marine systems in which denitrification occurs can be organized along a continuum ranging from (1) those in which nitrification and denitrification are tightly coupled in space and time to (2) those in which nitrate production and denitrification are relatively decoupled. In aquatic ecosystems, N inputs influence denitrification rates whereas hydrology and geomorphology influence the proportion of N inputs that are denitrified. Relationships between denitrification and water residence time and N load are remarkably similar across lakes, river reaches, estuaries, and continental shelves. Spatially distributed global models of denitrification suggest that continental shelf sediments account for the largest portion (44%) of total global denitrification, followed by terrestrial soils (22%) and oceanic oxygen minimum zones (OMZs; 14%). Freshwater systems (groundwater, lakes, rivers) account for about 20% and estuaries 1% of total global denitrification. Denitrification of land-based N sources is distributed somewhat differently. Within watersheds, the amount of land-based N denitrified is generally highest in terrestrial soils, with progressively smaller amounts denitrified in groundwater, rivers, lakes and reservoirs, and estuaries. A number of regional exceptions to this general trend of decreasing denitrification in a downstream direction exist, including significant denitrification in continental shelves of N from terrestrial sources. Though terrestrial soils and groundwater are responsible for much denitrification at the watershed scale, per-area denitrification rates in soils and groundwater (kg N x km(-2) x yr(-1)) are, on average, approximately one-tenth the per-area rates of denitrification in lakes, rivers, estuaries, continental shelves, or OMZs. A number of potential approaches to increase denitrification on the landscape, and thus decrease N export to sensitive coastal systems exist. However, these have not generally been widely tested for their effectiveness at scales required to significantly reduce N export at the whole watershed scale.     

Macroinvertebrate production and food web energetics in an industrially contaminated stream.

This study examines secondary production and periphyton-invertebrate food web energetics at two sites in an industrially contaminated, nutrient-enriched stream. Secondary production data and data from the literature were used to calculate potential amounts of mercury transferred from periphyton to chironomid larvae and into terrestrial food webs with emerging adults. The nutritional quality of periphyton was characterized using energy content, chlorophyll a, protein, ash-free dry mass (AFDM), and percentage of organic matter. Chironomid larvae (Orthocladiinae: Cricotopus spp.) comprised 96% of all macroinvertebrates collected from stones at the two sites. Cricotopus production was extremely high: production was 59.5 g AFDM x m(-2) x yr(-1) at the site upstream of a 1-ha settling basin and 32.4 g AFDM x m(-2) x yr(-1) at the site below the basin. Apparent differences in annual secondary production were associated with reduced organic content (i.e., nutritional quality) of the periphyton matrix under different loading of total suspended solids. The periphyton matrix at both sites was contaminated with inorganic (Hg(II)) and methyl (MeHg) mercury. The amount of Hg(II) potentially ingested by Cricotopus was calculated to be 49 mg Hg(II) x m(-1) x yr(-1) at the upstream site and 19 mg Hg(II)x m(-2) x yr(-1) at the downstream site. Mercury ingestion by Cricotopus at the downstream site was calculated to be 2% of the estimated annual deposition of particulate-bound Hg(II) to the stream bed. Emergence of adult Cricotopus was calculated to remove 563 microg Hg(II)x m(-2) x yr(-1) from the stream at the upstream site and 117 microg Hg(II) x m(-2) x yr(-1) at the downstream site, which amounted to 4.1 g Hg(II)/yr for the 2.1-km reach of stream included in this study. The ratio of metal export in emergence production to surface area for the study stream was 10 to 10(3) times higher than ratios calculated for lakes using data from the literature. This study is the first well-documented example of extremely high aquatic insect production in an industrially contaminated, nutrient-enriched stream, and it highlights the application of production measurements to examine the role of aquatic insect production in the trophic transfer of energy and persistent contaminants in aquatic food webs and into terrestrial food webs.     

Soil C and N changes with afforestation of grasslands across gradients of precipitation and plantation age

Afforestation, the conversion of unforested lands to forests, is a tool for sequestering anthropogenic carbon dioxide into plant biomass. However, in addition to altering biomass, afforestation can have substantial effects on soil organic carbon (SOC) pools, some of which have much longer turnover times than plant biomass. An increasing body of evidence suggests that the effect of afforestation on SOC may depend on mean annual precipitation (MAP). The goal of this study was to test how labile and bulk pools of SOC and total soil nitrogen (TN) change with afforestation across a rainfall gradient of 600-1500 mm in the Rio de la Plata grasslands of Argentina and Uruguay. The sites were all former grasslands planted with Eucalyptus spp. Overall, we found that afforestation increased (up to 1012 kg C x ha(-1) x yr(-1)) or decreased (as much as 1294 kg C x ha(-1) x yr(-1)) SOC pools in this region and that these changes were significantly related to MAP. Drier sites gained, and wetter sites lost, SOC and TN (r2 = 0.59, P = 0.003; and r2 = 0.57, P = 0.004, respectively). Labile C and N in microbial biomass and extractable soil pools followed similar patterns to bulk SOC and TN. Interestingly, drier sites gained more SOC and TN as plantations aged, while losses reversed as plantations aged in wet sites, suggesting that plantation age in addition to precipitation is a critical driver of changes in soil organic matter with afforestation. This new evidence implies that longer intervals between harvests for plantations could improve SOC storage, ameliorating the negative trends found in humid sites. Our results suggest that the value of afforestation as a carbon sequestration tool should be considered in the context of precipitation and age of the forest stand.     

Denitrification and the nitrogen budget of a reservoir in an agricultural landscape.

Denitrification is an important process in aquatic sediments, but its role has not been assessed in the N mass balance of upper-Midwestern (USA) reservoirs that receive large agricultural riverine N inputs. We used a 4400-ha reservoir to determine the role of denitrification in the N mass balance and effectiveness in reducing downstream transport of NO(3-)N. Sediment denitrification was (1) measured monthly (March 2002-March 2003) at eight sites in the Lake Shelbyville reservoir in central Illinois using the acetylene inhibition, chloramphenicol technique, (2) scaled to the overall reservoir and compared to N not accounted for in a mass balance, and (3) estimated indirectly using long-term (1981-2003) mass balances of N in the reservoir. Denitrification rates in the reservoir were high during spring and early summer of 2002, when maximum NO(3-)N concentrations were measured (10-14 mg NO(3-)N/L). We estimated that denitrification for the year was between 2580 and 5150 Mg N. Missing N from the mass balance was 3004 Mg N, suggesting that sediment denitrification was the sink. Areal rates of sediment denitrification in the reservoir ranged from 62 to 225 g N x m(-2) x yr(-1), with rates a function of both denitrification intensity (microg N x g dry mass x h(-1)) and the overall mass of sediment present. From 1981 to 2003 the average NO(3-)N inlet flux was 8900 Mg N/yr. About 58% of the total NO(3-)N input was removed, and annual NO(3-)N removed as a percentage of inputs was significantly related to reservoir retention time (average = 0.36 yr for the 23 years, range = 0.21-0.84 yr). By scaling denitrification in Lake Shelbyville to other reservoirs in Illinois, we estimated a sink of 48900 Mg N/yr. When combined with estimated in-stream denitrification, 60900 Mg N/yr was estimated to be removed by sediment denitrification. This reduces riverine export from Illinois to the Gulf of Mexico, where the flux during the 1990s was about 244000 Mg N/yr, and illustrates the importance of reservoir denitrification as an N sink in Midwestern agricultural landscapes.     

Impact of nutrient removal through harvesting on the sustainability of the boreal forest

The cycling of base cations (K, Ca, Mg, and Na) was investigated in a boreal balsam fir forest (the Lake Laflamme Watershed) between 1999 and 2005. Base cation budgets were calculated for the soil rooting zone that included atmospheric deposition and soil leaching losses, two scenarios of tree uptake (whole-tree and stem-only harvesting), and three scenarios of mineral weathering, leading to six different scenarios. In every scenario there was a net accumulation of Mg within the soil exchangeable reservoir, while Ca accumulated in four scenarios. Potassium was lost in five of the six scenarios. Contrary to Ca and Mg, immobilization of K within tree biomass (69 mol x ha(-1) x yr(-1)) was the main pathway of K losses from the soil exchangeable reservoir, being five times higher than losses via soil leaching (14 mol x ha(-1) x yr(-1)). The amounts of K contained within the aboveground biomass and the exchangeable soil reservoir were 3.3 kmol/ha and 4.2 kmol/ha, respectively. Whole-tree harvesting may thus remove 44% of the K that is readily available for cycling in the short term, making this forest sensitive to commercial forestry operations. Similar values of annual K uptake as well as a similar distribution of K between tree biomass and soil exchangeable reservoirs at 14 other coniferous sites, distributed throughout the boreal forest of Quebec, suggest that the Lake Laflamme Watershed results can be extrapolated to a much larger area. Stem-only harvesting, which would reduce K exports due to biomass removal by 60%, should be used for these types of forest.     

Necromass production: studies in undisturbed and logged Amazon forests.

Necromass stocks account for up to 20% of carbon stored in tropical forests and have been estimated to be 14-19% of the annual aboveground carbon flux. Both stocks and fluxes of necromass are infrequently measured. In this study, we directly measured the production of fallen coarse necromass (> or = 2 cm diameter) during 4.5 years using repeated surveys in undisturbed forest areas and in forests subjected to reduced-impact logging at the Tapajos National Forest, Belterra, Brazil (3.08 degrees S, 54.94 degrees W). We also measured fallen coarse necromass and standing dead stocks at two times during our study. The mean (SE) annual flux into the fallen coarse necromass pool in undisturbed forest of 6.7 (0.8) Mg x ha(-1) x yr(-1) was not significantly different from the flux under a reduced-impact logging of 8.5 (1.3) Mg x ha(-1) x yr(-1) With the assumption of steady state, the instantaneous decomposition constants for fallen necromass in undisturbed forests were 0.12 yr(-1) for large, 0.33 yr(-1) for medium, and 0.47 yr(-1) for small size classes. The mass weighted decomposition constant was 0.15 yr(-1) for all fallen coarse necromass. Standing dead wood had a residence time of 4.2 years, and approximately 0.9 Mg x ha(-1) x yr(-1) of this pool was respired annually to the atmosphere through decomposition. Coarse necromass decomposition at our study site accounted for 12% of total carbon remineralization, and total aboveground coarse necromass was 14% of the aboveground biomass. Use of mortality rates to calculate production of coarse necromass leads to an underestimation of coarse necromass production by 45%, suggesting that nonlethal disturbance such as branch fall contributes significantly to this flux. Coarse necromass production is an important component of the tropical forest carbon cycle that has been neglected in most previous studies or erroneously estimated.     

Flow of chemical energy in Alwar jheel of Yamuna basin near Allahabad

The water quality, rate of energy transformation, chemical composition of producers and flow of chemical energy were studied in both feeding river Yamuna and Alwar jheel near Allahabad. As the river Yamuna had high value of alkalinity (210.0 mgl(-1)), conductance (518.0 micromhos), dissolved solids (260.0 mgl(-1)), hardness (162.0 mgl(-1)) and chloride (54.6 mgl(-1)) jheel also showed high values of these parameters. The rate of energy transformation from kinetic radiant energy to chemical energy was very high in the jheel 32,315 Cal m(-2) day(-1) of which 25,620 Cal m(-2) day(-1) was contributed by aquatic plants. Out of 11,764 x 10(4) Kcal ha(-1) yr(-1) total energy fixed in the system, producers stored 7,154 x 10(4) Kcal ha(-1) yr(-1) and the rest was lost as heat of respiration. The pattern of storage of energy was different in two groups of producers and thus most of the energy fixed by phytoplankton was stored as protein (56.2%) and less as carbohydrate (11.7%) while aquatic plants stored more energy as carbohydrate (40.8%) than protein (23.2%). The chemical energy obtained from the system was 1,85,000 Kcal ha(-1) yr(-1) and thus only 0.260% of the chemical energy stored by producers was harvested. The potential chemical energy resource in the jheel was 81.4 x 10(4) Kcal ha(-1) yr(-1) of which only 22.6% was harvested in the jheel and there is enough scope for further enhancement.