Dissolved and particulate nutrient flux from three adjacent agricultural watersheds: A five-year study

Fluxes of dissolved and particulate nitrogen (N) and phosphorus(P) from three adjacent watersheds were quantified with ahigh-resolution sampling program over a five-year period. The watershedsvary by an order of magnitude in area (12,875, 7968 and 1206 ha), and inall three watersheds intensive agriculture comprises > 90% ofland. Annual fluxes of dissolved N and P per unit watershed area (exportcoefficients) varied 2X among watersheds, and patterns were notdirectly related to watershed size. Over the five-year period, meanannual flux of soluble reactive P (SRP) was 0.583 kg P ·ha^–1 · yr^–1 from the smallestwatershed and 0.295 kg P · ha^–1 ·yr^–1 from the intermediate-sized watershed, which hadthe lowest SRP flux. Mean annual flux of nitrate was 20.53 kg N ·ha^–1 · yr^–1 in the smallestwatershed and 44.77 kg N · ha^–1 ·yr^–1 in the intermediate-sized watershed, which had thehighest nitrate flux. As a consequence, the export ratio of dissolvedinorganic N to SRP varied from 80 (molar) in the smallest watershed to335 in the intermediate-sized watershed. Because most N was exported asnitrate, differences among watersheds in total N flux were similar tothose for nitrate. Hence, the total N:P export ratio was 42(molar) for the smallest watershed and 109 for the intermediate-sizedwatershed. In contrast, there were no clear differences among watershedsin the export coefficients of particulate N, P, or carbon, even though> 50% of total P was exported as particulate P in allwatersheds. All nutrient fractions were exported at higher rates in wetyears than in dry years, but precipitation-driven variability in exportcoefficients was greater for particulate fractions than for dissolvedfractions.Examination of hydrological regimes showed that, for all nutrientfractions, most export occurred during stormflow. However, theproportion of nitrate flux exported as baseflow was much greater thanthe proportion of SRP flux exported as baseflow, for all threewatersheds (25–37% of nitrate exported as baseflow vs.3–13% of SRP exported as baseflow). In addition, baseflowcomprised a greater proportion of total discharge in theintermediate-sized watershed (43.7% of total discharge) than theother two watersheds (29.3 and 30.1%). Thus, higher nitrateexport coefficients in the intermediate-sized watershed may haveresulted from the greater contribution of baseflow in this watershed.Other factors potentially contributing to higher nitrate exportcoefficients in this watershed may be a thicker layer of loess soils anda lower proportion of riparian forest than the other watersheds. Theamong-watershed variability in SRP concentrations and exportcoefficients remains largely unexplained, and might represent theminimum expected variation among similar agriculturalwatersheds.     

Does Anthropogenic Nitrogen Enrichment Increase Organic Nitrogen Concentrations in Runoff from Forested and Human-dominated Watersheds?

Although the effects of anthropogenic nitrogen (N) inputs on the dynamics of inorganic N in watersheds have been studied extensively, “the influence of N enrichment on organic N loss” is not as well understood. We compiled and synthesized data on surface water N concentrations from 348 forested and human-dominated watersheds with a range of N loads (from less than 100 to 7,100 kg N km⁻² y⁻¹) to evaluate the effects of N loading via atmospheric deposition, fertilization, and wastewater on dissolved organic N (DON) concentrations. Our results indicate that, on average, DON accounts for half of the total dissolved N (TDN) concentrations from forested watersheds, but it accounts for a smaller fraction of TDN in runoff from urban and agricultural watersheds with higher N loading. A significant but weak correlation (r ² = 0.06) suggests that N loading has little influence on DON concentrations in forested watersheds. This result contrasts with observations from some plot-scale N fertilization studies and suggests that variability in watershed characteristics and climate among forested watersheds may be a more important control on DON losses than N loading from atmospheric sources. Mean DON concentrations were positively correlated, however, with N load across the entire land-use gradient (r ² = 0.37, P < 0.01), with the highest concentrations found in agricultural and urban watersheds. We hypothesize that both direct contributions of DON from wastewater and agricultural amendments and indirect transformations of inorganic N to organic N represent important sources of DON to surface waters in human-dominated watersheds. We conclude that DON is an important component of N loss in surface waters draining forested and human-dominated watersheds and suggest several research priorities that may be useful in elucidating the role of N enrichment in watershed DON dynamics.     

Urban influences on the nitrogen cycle in Puerto Rico

Anthropogenic actions are altering fluxes of nitrogen (N) in the biosphere at unprecedented rates. Efforts to study these impacts have concentrated in the Northern hemisphere, where experimental data are available. In tropical developing countries, however, experimental studies are lacking. This paper summarizes available data and assesses the impacts of human activities on N fluxes in Puerto Rico, a densely populated Caribbean island that has experienced drastic landscape transformations over the last century associated with rapid socioeconomic changes. N yield calculations conducted in several watersheds of different anthropogenic influences revealed that disturbed watersheds export more N per unit area than undisturbed forested watersheds. Export of N from urban watersheds ranged from 4.8 kg ha^?1 year^?1 in the Río Bayamón watershed to 32.9 kg ha^?1 year^?1 in the highly urbanized Río Piedras watershed and 33.3 kg ha^?1 year^?1 in the rural-agricultural Río Grande de Añasco watershed. Along with land use, mean annual runoff explained most of the variance in fluvial N yield. Wastewater generated in the San Juan Metropolitan Area receives primary treatment before it is discharged into the Atlantic Ocean. These discharges are N-rich and export large amounts of N to the ocean at a rate of about 140 kg ha^?1 year^?1. Data on wet deposition of inorganic N ( $\hbox{NH}_{4}^{+}+\hbox{NO}_{3}^{-}Anthropogenic actions are altering fluxes of nitrogen (N) in the biosphere at unprecedented rates. Efforts to study these impacts have concentrated in the Northern hemisphere, where experimental data are available. In tropical developing countries, however, experimental studies are lacking. This paper summarizes available data and assesses the impacts of human activities on N fluxes in Puerto Rico, a densely populated Caribbean island that has experienced drastic landscape transformations over the last century associated with rapid socioeconomic changes. N yield calculations conducted in several watersheds of different anthropogenic influences revealed that disturbed watersheds export more N per unit area than undisturbed forested watersheds. Export of N from urban watersheds ranged from 4.8 kg ha⁻¹ year⁻¹ in the Río Bayamón watershed to 32.9 kg ha⁻¹ year⁻¹ in the highly urbanized Río Piedras watershed and 33.3 kg ha⁻¹ year⁻¹ in the rural-agricultural Río Grande de A?asco watershed. Along with land use, mean annual runoff explained most of the variance in fluvial N yield. Wastewater generated in the San Juan Metropolitan Area receives primary treatment before it is discharged into the Atlantic Ocean. These discharges are N-rich and export large amounts of N to the ocean at a rate of about 140 kg ha⁻¹ year⁻¹. Data on wet deposition of inorganic N () suggest that rates of atmospheric N deposition are increasing in the pristine forests of Puerto Rico. Stationary and mobile sources of NO _x (NO+NO₂) and N₂O generated in the large urban centers may be responsible for this trend. Comprehensive measurements are required in Puerto Rico to quantitatively characterize the local N cycle. More research is required to assess rates of atmospheric N deposition, N fixation in natural and human-dominated landscapes, N-balance associated with food and feed trade, and denitrification.     

Forest nitrogen sinks in large eastern U.S. watersheds: estimates from forest inventory and an ecosystem model

The eastern U.S. receives elevated rates of Ndeposition compared to preindustrial times, yetrelatively little of this N is exported indrainage waters. Net uptake of N into forestbiomass and soils could account for asubstantial portion of the difference between Ndeposition and solution exports. We quantifiedforest N sinks in biomass accumulation andharvest export for 16 large river basins in theeastern U.S. with two separate approaches: (1)using growth data from the USDA ForestService's Forest Inventory and Analysis (FIA)program, and (2) using a model of forestnitrogen cycling (PnET-CN) linked to FIAinformation on forest age-class structure. Themodel was also used to quantify N sinks in soiland dead wood, and nitrate losses below therooting zone. Both methods agreed that netgrowth rates were highest in the relativelyyoung forests on the Schuylkill watershed, andlowest in the cool forests of northern Maine. Across the 16 watersheds, wood export removedan average of 2.7 kg N ha^–1 yr^–1(range: 1–5 kg N ha^–1 yr^–1), andstanding stocks increased by 4.0 kg N ha^–1yr^–1 (–3 to 8 kg N ha^–1 yr^–1). Together, these sinks for N in woody biomassamounted to a mean of 6.7 kg N ha^–1yr^–1 (2–9 kg N ha^–1 yr^–1), or73% (15–115%) of atmospheric N deposition. Modeled rates of net N sinks in dead wood andsoil were small; soils were only a significantnet sink for N during simulations ofreforestation of degraded agricultural sites. Predicted losses of nitrate depended on thecombined effects of N deposition, and bothshort- and long-term effects of disturbance. Linking the model with forest inventoryinformation on age-class structure provided auseful step toward incorporating realisticpatterns of forest disturbance status acrossthe landscape.     

Changing Global Climate: Historical Carbon and Nitrogen Budgets and Projected Responses of Ohio’s Cropland Ecosystems

As the evidence of global climate change continues to mount, its consequences for cropland productivity assume particular significance. Against the backdrop of past agricultural practices, simulation models offer a glimpse into the future, showing the effect of temperature changes on crop production. In this study, we first quantified the carbon (C) and nitrogen (N) budgets of Ohios cropland ecosystems using inventory yield data of corn for grain, oat, and all wheat for the period 1866–1996 and soybean for the period 1924–96. Then we explored the responses of Ohios continuous soybean croplands to changes in temperature, carbon dioxide (CO₂) concentration, initial soil organic C and N (SOC-N) pools, soil texture, and management practices by developing a simple cropland ecosystem model (CEM) and performing a long-term sensitivity analysis. Finally, CEM simulations were evaluated against independent observations of SOC values (0–19 cm) averaged over 470 northwest Ohio sites between 1954 and 1987 under conventional tillage and rotations of corn–soybean–winter wheat by using the historical yield data (r ² = 0.8). The C contents per hectare of crop harvests increased by 178% for oats, 300% for corn for grain, and 652% for all wheat between 1866 and 1996 and by 305% for soybean between 1924 and 1996. Ohio croplands acted as C–N sources, releasing average net ecosystem emissions (NEE), including the removal of harvested C–N, of 4,598 kg CO₂ ha^–1 and 141 kg N ha^–1 in 1886 and 205 kg CO₂ ha^–1 (except for the corn-for-grain cropland) and 39 kg N ha^–1 in 1996. The continuous corn croplands continued to become a C sink, sequestering 255 kg C ha^–1 in 1996. Results of the sensitivity analysis for Ohios continuous soybean croplands revealed that the SOC pool increased by 6.9% and decreased by 7.5% in response to a doubled CO₂ concentration and a temperature increase of 2.8°C over 100 years, respectively. The sequestration potential of the SOC pool increased by 6.5% at a rate of 24.6 kg C ha^–1 y^–1 for the same period with finer soil texture (loam to silty clay loam). The shift from conventional to conservation residue practice led to an 11% increase in the steady-state SOC storage at a rate of 42 kg C ha^–1 y^–1 for 100 years.     

Biogeochemistry of unpolluted forested watersheds in the Oregon Cascades: temporal patterns of precipitation and stream nitrogen fluxes

We analyzed long-term organic and inorganic nitrogen inputs and outputs in precipitation and streamwater in six watersheds at the H.J. Andrews Experimental Forest in the central Cascade Mountains of Oregon. Total bulk N deposition, averaging 1.6 to 2.0 kg N ha^–1 yr^–1, is low compared to other sites in the United States and little influenced by anthropogenic N sources. Streamwater N export is also low, averaging <1 kg ha^–1 yr^–1. DON is the predominant form of N exported from all watersheds, followed by PON, NH₄-N, and NO₃-N. Total annual stream discharge was a positive predictor of annual DON output in all six watersheds, suggesting that DON export is related to regional precipitation. In contrast, annual discharge was a positive predictor of annual NO₃-N output in one watershed, annual NH₄-N output in three watersheds, and annual PON output in three watersheds. Of the four forms of N, only DON had consistent seasonal concentration patterns in all watersheds. Peak streamwater DON concentrations occurred in November-December after the onset of fall rains but before the peak in the hydrograph, probably due to flushing of products of decomposition that had built up during the dry summer. Multiple biotic controls on the more labile nitrate and ammonium concentrations in streams may obscure temporal DIN flux patterns from the terrestrial environment. Results from this study underscore the value of using several watersheds from a single climatic zone to make inferences about controls on stream N chemistry; analysis of a single watershed may preclude identification of geographically extensive mechanisms controlling N dynamics.     

Suppression of nitrate formation within an exotic conifer plantation

Summary Nitrate-N losses to stream waters and soil inorganic N pools, nitrifying potentials and NO₃-N production rates were measured in 2 adjacent watersheds, one used as pasture and the other planted in exotic conifer forest (Pinus radiata D. Don). Estimated NO₃-N loss to stream waters draining the pine and pasture watersheds were 0.6kg ha⁻¹ y⁻¹ and 7.6 kg ha⁻¹ y⁻¹ respectively. Ammonium-N pool sizes were not significantly different between soils in the two watersheds but NO₃−N pools and nitrifying potentials were always lower in the pine watershed soil samples. Laboratory incubation experiments indicated that suppression of NO₃−N formation in pine watershed soils required the presence of live tree roots and was not due to the direct action of allelopathic chemicals on nitrifiers.     

Yield of nitrogen from minimally disturbed watersheds of the United States

Watersheds of the US Geological Survey's Hydrologic Benchmark Network program were used in estimating annual yield of total nitrogen and nitrogen fractions (ammonium, nitrate, dissolved organic N, particulate N) in relation to amount of runoff, elevation, and watershed area. Only watersheds minimally disturbed with respect to the nitrogen cycle were used in the analysis (mostly natural vegetation cover, no point sources of N, atmospheric deposition of inorganic N < 10 kg ha^–1 y^–1). Statistical analysis of the yields of total nitrogen and nitrogen fractions showed that elevation and watershed area bear no significant relationship to nitrogen yield for these watersheds. The yields of total nitrogen and nitrogen fractions are, however, strongly related to runoff (r ² = 0.91 for total N). Annual yield increases as runoff increases, but at a rate lower than runoff; annual discharge-weighted mean concentrations decline as annual runoff increases. Yields of total nitrogen and most nitrogen fractions bear a relationship to runoff that is nearly indistinguishable from a relationship that was documented previously for minimally disturbed watersheds of the American tropics. Overall, the results suggest strong interlatitudinal convergence of yields and percent fractionation for nitrogen in relation to runoff.     

Nitrogen Export from Forested Watersheds in the Oregon Coast Range: The Role of N<Subscript>2</Subscript>-fixing Red Alder

Variations in plant community composition across the landscape can influence nutrient retention and loss at the watershed scale. A striking example of plant species importance is the influence of N₂-fixing red alder (Alnus rubra) on nutrient cycling in the forests of the Pacific Northwest. To understand the influence of red alder on watershed nutrient export, we studied the chemistry of 26 small watershed streams within the Salmon River basin of the Oregon Coast Range. Nitrate and dissolved organic nitrogen (DON) concentrations were positively related to broadleaf cover (dominated by red alder: 94% of basal area), particularly when near-coastal sites were excluded (r ² = 0.65 and 0.68 for nitrate-N and DON, respectively). Nitrate and DON concentrations were more strongly related to broadleaf cover within entire watersheds than broadleaf cover within the riparian area alone, which indicates that leaching from upland alder stands plays an important role in watershed nitrogen (N) export. Nitrate dominated over DON in hydrologic export (92% of total dissolved N), and nitrate and DON concentrations were strongly correlated. Annual N export was highly variable among watersheds (2.4–30.8 kg N ha^–1 y^–1), described by a multiple linear regression combining broadleaf and mixed broadleaf–conifer cover (r² = 0.74). Base cation concentrations were positively related to nitrate concentrations, which suggests that nitrate leaching increases cation losses. Our findings provide evidence for strong control of ecosystem function by a single plant species, where leaching from N saturated red alder stands is a major control on N export from these coastal watersheds.     

Long-Term Trends in Stream Nitrate Concentrations and Losses Across Watersheds Undergoing Recovery from Acidification in the Czech Republic

Stream nitrogen (N) export and nitrate concentration were measured at 14 forested watersheds (GEOMON network) in the Czech Republic between 1994 and 2005. In the last several decades, emissions of sulfur (S) and N compounds have decreased throughout much of Europe. In the Czech Republic, atmospheric deposition of S has decreased substantially since the beginning of 1990s, whereas N deposition remains largely unchanged at most sites. The mean dissolved inorganic nitrogen (DIN) streamwater export ranged from 0.2 to 12.2 kg ha⁻¹ y⁻¹ at the GEOMON sites. Despite decades of elevated N deposition, 44–98% of DIN inputs to these watersheds were retained or denitrified, and many watersheds showed seasonal variation in nitrate concentrations. Dissolved organic N export was quantified in 1 year only and ranged from 0.05 to 3.5 kg ha⁻¹ y⁻¹. Spatial variability in DIN export among watersheds was best explained by spatial variability in average acidic deposition, particularly S deposition (R ² = 0.81, P < 0.001); DIN input and forest floor carbon:nitrogen (C/N) also provided significant explanatory power. DIN export was strongly influenced by the forest floor C/N ratio and depth of the forest floor soils (R ² = 0.72, P < 0.001). The only variable that predicted variations in forest floor C/N (R ² = 0.32, P < 0.05) among watersheds was S deposition. Forest floor depth was also related to deposition variables, with S deposition providing the most explanatory power (R ² = 0.50, P < 0.01). Variation in forest floor depth was also associated with climatic factors (precipitation and temperature). Temporal variability in DIN export was primarily associated with changes in acidic deposition over time; S deposition explained 41% of variability in DIN exports among all watersheds and years. Extensive acidification of forested watersheds was associated with the extraordinarily high S inputs to much of the Czech Republic during earlier decades. We hypothesize that recovery from acidification has led to improved tree health as well as enhanced microbial activity in the forest floor. As these watersheds move into a new regime with dramatically lower sulfur inputs, we expect continued declines in nitrate output.     

Transformation and retention of nitrogen in a coastal forest ecosystem

Transformations and fluxes of N were examined in three forested sites located along a gradient of soil texture in the coastal forests of the Waquoit Bay watershed on Cape Cod. Total N leaching losses to ground water were 0.5 kg ha^-1 yr^-1 in the loamy sand site and 1.5 kg ha^-1 yr^-1 in the fine sand site. Leaching loss to groundwater was not measured in the coarse sand site due to the prohibitive depth of the water table but total N leaching loss to 1m depth in the mineral soil was 3.9 kg ha^-1 yr^-1. DON accounted for most of the leaching losses below the rooting zone (77–89%) and through the soil profile to ground water (60%–80%). Differences in DON retention capacity of the mineral soil in the sites along the soil texture gradient were most likely related to changes in mineral soil particle surface area and percolation rates associated with soil texture. Forests of the watershed functioned as a sink for inorganic N deposited on the surface of the watershed in wet and dry deposition but a source of dissolved organic N to ground water and adjoining coastal ecosystems.     

Regional analysis of inorganic nitrogen yield and retention in high-elevation ecosystems of the Sierra Nevada and Rocky Mountains

Yields and retention of dissolved inorganic nitrogen (DIN: NO₃ ^– + NH₄ ⁺) and nitrate concentrations in surface runoff are summarized for 28 high elevation watersheds in the Sierra Nevada of California and Rocky Mountains of Wyoming and Colorado. Catchments ranged in elevation from 2475 to 3603 m and from 15 to 1908 ha in area. Soil cover varied from 5% to nearly 97% of total catchment area. Runoff from these snow-dominated catchments ranged from 315 to 1265 mm per year. In the Sierra Nevada, annual volume-weighted mean (AVWM) nitrate concentrations ranged from 0.5 to 13 M (overall average 5.4 M), and peak concentrations measured during snowmelt ranged from 1.0 to 38 M. Nitrate levels in the Rocky Mountain watersheds were about twice those in the Sierra Nevada; average AVWM NO₃ ^– was 9.4 M and snowmelt peaks ranged from 15 to 50 M. Mean DIN loading to Rocky Mountain watersheds, 3.6 kg ha^–1 yr^–1, was double the average measured for Sierra Nevada watersheds, 1.8 kg ha^–1 yr^–1. DIN yield in the Sierra Nevada, 0.69 kg ha^–1 yr^–1, was about 60% that measured in the Rocky Mountains, 1.1 kg ha^–1 yr^–1. Net inorganic N retention in Sierra Nevada catchments was 1.2 kg ha^–1 yr^–1 and represented about 55% of annual DIN loading. DIN retention in the Rocky Mountain catchments was greater in absolute terms, 2.5 kg ha^–1 yr^–1, and as a percentage of DIN loading, 72%.A correlation analysis using DIN yield, DIN retention and surface water nitrate concentrations as dependent variables and eight environmental features (catchment elevation, slope, aspect, roughness, area, runoff, soil cover and DIN loading) as independent variables was conducted. For the Sierra Nevada, elevation and soil cover had significant (p > 0.1) Pearson product moment correlations with catchment DIN yield, AVWM and peak snowmelt nitrate concentrations and DIN retention rates. Log-linear regression models using soil cover as the independent variable explained 82% of the variation in catchment DIN retention, 92% of the variability in AVWM nitrate and 85% of snowmelt peak NO₃ ^–. In the Rocky Mountains, soil cover was significantly (p < 0.05) correlated with DIN yield, AVWM NO₃ ^– and DIN retention expressed as a percentage of DIN loading (%DIN retention). Catchment mean slope and terrain roughness were positively correlated with steam nitrate concentrations and negatively related to %DIN retention. About 91% of the variation in DIN yield and 79% of the variability in AVWM NO₃ ^– were explained by log-linear models based on soil cover. A log-linear regression based on soil cover explained 90% of the variation of %DIN retention in the Rocky Mountains.     

Retention of soluble organic nutrients by a forested ecosystem

We document an example of a forested watershed at the Coweeta HydrologicLaboratory with an extraordinary tendency to retain dissolved organic matter(DOM) generated in large quantities within the ecosystem. Our objectives weretodetermine fluxes of dissolved organic C, N, and P (DOC, DON, DOP,respectively),in water draining through each stratum of the ecosystem and synthesizeinformation on the physicochemical, biological and hydrologic factors leadingtoretention of dissolved organic nutrients in this ecosystem. The ecosystemretained 99.3, 97.3, and 99.0% of water soluble organic C, N and P,respectively, produced in litterfall, throughfall, and root exudates. Exportsinstreamwater were 4.1 kg ha^–1yr^–1of DOC, 0.191 kg ha^–1 yr^–1 ofDON, and 0.011 kg ha^–1 yr^–1 ofDOP. Fluxes of DON were greater than those of inorganic N in all strata. MostDOC, DON, and DOP was removed from solution in the A and B horizons, with DOCbeing rapidly adsorbed to Fe and Al oxyhydroxides, most likely by ligandexchange. DON and DOC were released gradually from the forest floor over theyear. Water soluble organic C produced in litterfall and throughfall had adisjoint distribution of half-decay times with very labile and veryrefractory fractions so that most labile DOC was decomposed before beingleachedinto the mineral soil and refractory fractions dominated the DOC transportedthrough the ecosystem. We hypothesize that this watershed retained solubleorganic nutrients to an extraordinary degree because the soils have very highcontents of Fe and Al oxyhydroxides with high adsorption capacities and becausethe predominant hydrologic pathway is downwards as unsaturated flow through astrongly adsorbing A and B horizon. The well recognized retention mechanismsforinorganic nutrients combine with adsorption of DOM and hydrologic pathway toefficiently prevent leaching of both soluble inorganic andorganic nutrients in this watershed.     

Nitrogen mineralisation in podzol soils under boreal Scots pine and Norway spruce stands

N mineralisation was investigated in the mor humus layer of a podzol at a forested catchment area of Saarejärve Lake in Eastern Estonia. The investigated areas were pine (Rhodococcumunderstorey) and spruce (Vaccinium understorey) stands, which are permanent sample plots of an integrated monitoring network. The seasonal pattern of net N mineralisation was studied by incubating undisturbed cores of mor humus (0–8 cm) in buried polyethylene bags in situ. Samples were collected and incubated between July 1996 and April 1998. The period of incubation was approximately 1 month, except for wintertime when incubation lasted till thawing of ground (5 months). The amounts of mineral nitrogen formed during monthly incubations in vegetation period vary considerably (0.4–8.7 kg ha^–1). About 70% of the variation of net ammonification could be explained by environmental factors - temperature, initial moisture and pH. Ammonium was the dominant form of mineral nitrogen, which is typical for mor humus. The rate of nitrification was very low, and most of the annual net nitrification occurred during just one or two months (May–June, October) depending on site and year. Measured annual net N mineralisation was 29.2 kg ha^–1 for the spruce stand and 23.6 kg ha^–1 for the pine stand. These measures were found to be in good accordance with other N-fluxes in the ecosystem.     

N deposition, N transformation and N leaching in acid forest soils

Nitrogen deposition, mineralisation, uptake and leaching were measured on a monthly basis in the field during 2 years in six forested stands on acidic soils under mountainous climate. Studies were conducted in three Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] plantations (D20: 20 year; D40: 40 yr; D60: 60 yr) on abandoned croplands in the Beaujolais Mounts; and two spruce (Picea abies Karst.) plantations (S45: 45 yr; S90: 90 yr) and an old beech (Fagus sylvatica L.) stand (B150: 150 yr) on ancient forest soils in a small catchment in the Vosges Mountains. N deposition in throughfall varied between 7–8 kg ha^–1 year^–1 (D20, B150, S45) and 15–21 kg ha^–1 yr^–1 (S90, D40, D60). N in annual litterfall varied between 20–29 kg ha^–1 (D40, D60, S90), and 36–43 kg ha^–1 (D20, S45, B150). N leaching below root depth varied among stands within a much larger range, between 1–9 kg ha^–1 yr^–1 (B150, S45, D60) and 28–66 kg ha^–1 yr^–1 (D40, S90, D20), with no simple relationship with N deposition, or N deposition minus N storage in stand biomass. N mineralisation was between 57–121 kg ha^–1 yr^–1 (S45, D40, S90) and between 176–209 kg ha^–1 yr^–1 in (B150, D60 and D20). The amounts of nitrogen annually mineralised and nitrified were positively related. Neither general soil parameters, such as pH, soil type, base saturation and C:N ratio, nor deposition in throughfall or litterfall were simply related to the intensity of mineralisation and/or nitrification. When root uptake was not allowed, nitrate leaching increased by 11 kg ha^–1 yr^–1 at S45, 36 kg ha^–1 yr^–1 at S90 and between 69 and 91 kg ha^–1 yr^–1 at D20, D40, B150 and D60, in relation to the nitrification rates of each plot. From this data set and recent data from the literature, we suggest that: high nitrification and nitrate leaching in Douglas-fir soils was likely related to the former agricultural land use. High nitrification rate but very low nitrate leaching in the old beech soil was related to intense recycling of mineralised N by beech roots. Medium nitrification and nitrate leaching in the old spruce stand was related to the average level of N deposition and to the deposition and declining health of the stand. Very low nitrification and N leaching in the young spruce stand were considered representative of fast growing spruce plantations receiving low N deposition on acidic soils of ancient coniferous forests. Consequently, we suggest that past land use and fine root cycling (which is dependent on to tree species and health) should be taken into account to explain the variability in the relation between N deposition and leaching in forests.     

Atmospheric deposition and solute export in giant sequoia — mixed conifer watersheds in the Sierra Nevada,California

Atmospheric depostion and stream discharge and solutes were measured for three years (September 1984 — August 1987) in two mixed conifer watersheds in Sequoia National Park, in the southern Sierra Nevada of California. The Log Creek watershed (50 ha, 2067–2397 m elev.) is drained by a perennial stream, while Tharp's Creek watershed (13 ha, 2067–2255 m elev.) contains an intermittent stream. Dominant trees in the area include Abies concolor (white fir), Sequoiadendron giganteum (giant sequoia), A. magnifica (red fir), and Pinus lambertiana (sugar pine). Bedrock is predominantly granite and granodiorite, and the soils are mostly Pachic Xerumbrepts. Over the three year period, sulfate (SO₄ ^2–), nitrate (NO₃ ^–), and chloride (Cl^–1) were the major anions in bulk precipitation with volume-weighted average concentrations of 12.6, 12.3 and 10.0 eq/1, respectively. Annual inputs of NO₃-N, NH₄-N and SO₄-S from wet deposition were about 60 to 75% of those reported from bulk deposition collectors. Discharge from the two watersheds occurs primarily during spring snowmelt. Solute exports from Log and Tharp's Creeks were dominated by HCO₃ ^–, Ca²⁺ and Na⁺, while H⁺, NO₃ ^–, NH₄ ⁺ and PO₄ ^3– outputs were relatively small. Solute concentrations were weakly correlated with instantaneous stream flow for all solutes (r² <0.2) except HCO₃ ^– (Log Cr. r² = 0.72; Tharp's Cr. r² = 0.38), Na⁺ (Log Cr. r² = 0.56; Tharp's Cr. r² = 0.47), and silicate (Log Cr. r² = 0.71; Tharp's Cr. r² = 0.49). Mean annual atmospheric contributions of NO₃-N (1.6 kg ha^–1), NH₄-N (1.7 kg ha^–1), and SO₄-S (1.8 kg ha^–1), which are associated with acidic deposition, greatly exceed hydrologic losses. Annual watershed yields (expressed as eq ha^–1) of HCO₃ ^– exceeded by factors of 2.5 to 37 the annual atmospheric deposition of H⁺.     

Fine root growth phenology,production, and turnover in a northern hardwood forest ecosystem

A large part of the nutrient flux in deciduous forests is through fine root turnover, yet this process is seldom measured. As part of a nutrient cycling study, fine root dynamics were studied for two years at Huntington Forest in the Adirondack Mountain region of New York, USA. Root growth phenology was characterized using field rhizotrons, three methods were used to estimate fine root production, two methods were used to estimate fine root mortality, and decomposition was estimated using the buried bag technique. During both 1986 and 1987, fine root elongation began in early April, peaked during July and August, and nearly ceased by mid-October. Mean fine root ( 3 mm diameter) biomass in the surface 28-cm was 2.5 t ha^–1 and necromass was 2.9 t ha^–1. Annual decomposition rates ranged from 17 to 30% beneath the litter and 27 to 52% at a depth of 10 cm. Depending on the method used for estimation, fine root production ranged from 2.0 to 2.9 t ha^–1, mortality ranged from 1.8 to 3.7 t ha^–1 yr^–1, and decomposition was 0.9 t ha^–1 yr^–1. Thus, turnover ranged from 0.8 to 1.2 yr^–1. The nutrients that cycled through fine roots annually were 4.5–6.1 kg Ca, 1.1–1.4 kg Mg, 0.3–0.4 kg K, 1.2–1.7 kg P, 20.3–27.3 kg N, and 1.8–2.4 kg S ha^–1. Fine root turnover was less important than leaf litterfall in the cycling of Ca and Mg and was similar to leaf litterfall in the amount of N, P, K and S cycled.     

Nitrogen yields from undisturbed watersheds in the Americas

Yields of total fixed nitrogen and nitrogen fractions are summarized for thirty-one watersheds in which anthropogenic disturbance of the nitrogen cycle, either through land use or atmospheric deposition, is negligible or slight. These yields are taken as representative of background conditions over a broad range of watershed areas, elevations, and vegetation types. The data set focuses on watersheds of the American tropics, but also includes information on the Gambia River (Africa) and some small watersheds in the Sierra Nevada of California. For the tropical watersheds, total nitrogen yield averages 5.1 kg ha^–1 y^–1. On average, 30% of the total is particulate and 70% is dissolved. Of the dissolved fraction, an average of 50% is organic and 50% is inorganic, of which 20% is ammonium and 80% is nitrate. Yields are substantially lower than previously estimated for background conditions. Yields of all nitrogen fractions are strongly related to runoff, which also explains a large percentage of variance in yield of total nitrogen (r²=0.85). For total nitrogen and nitrogen fractions, yield increases at about two-thirds the rate of runoff; concentration decreases as runoff increases. There is a secondary but significant positive relationship between elevation and yield of DIN. Ratios DON/TDN and PN/TN both are related to watershed area rather than runoff; DON/TDN decreases and PN/TN increases toward higher stream orders. The analysis suggests for tropical watersheds the existence of mechanisms promoting strong homeostasis in the yield of N and its fractions for a given moisture regime, as well as predictable downstream change in proportionate representation N fractions. Yields and concentrations for small tropical watersheds are much larger than for the few temperate ones with which comparisons are possible.     

The Role of Dissolved Organic Carbon, Dissolved Organic Nitrogen, and Dissolved Inorganic Nitrogen in a Tropical Wet Forest Ecosystem

Although tropical wet forests play an important role in the global carbon (C) and nitrogen (N) cycles, little is known about the origin, composition, and fate of dissolved organic C (DOC) and N (DON) in these ecosystems. We quantified and characterized fluxes of DOC, DON, and dissolved inorganic N (DIN) in throughfall, litter leachate, and soil solution of an old-growth tropical wet forest to assess their contribution to C stabilization (DOC) and to N export (DON and DIN) from this ecosystem. We found that the forest canopy was a major source of DOC (232 kg C ha^–1 y^–1). Dissolved organic C fluxes decreased with soil depth from 277 kg C ha^–1 y^–1 below the litter layer to around 50 kg C kg C ha^–1 y^–1 between 0.75 and 3.5m depth. Laboratory experiments to quantify biodegradable DOC and DON and to estimate the DOC sorption capacity of the soil, combined with chemical analyses of DOC, revealed that sorption was the dominant process controlling the observed DOC profiles in the soil. This sorption of DOC by the soil matrix has probably led to large soil organic C stores, especially below the rooting zone. Dissolved N fluxes in all strata were dominated by mineral N (mainly NO₃⁻). The dominance of NO₃^– relative to the total amount nitrate of N leaching from the soil shows that NO₃^– is dominant not only in forest ecosystems receiving large anthropogenic nitrogen inputs but also in this old-growth forest ecosystem, which is not N-limited.     

Production,standing biomass and natural abundance of <Superscript>15</Superscript>N and <Superscript>13</Superscript>C in ectomycorrhizal mycelia collected at different soil depths in two forest types

Nutrient uptake by forest trees is dependent on ectomycorrhizal (EM) mycelia that grow out into the soil from the mycorrhizal root tips. We estimated the production of EM mycelia in root free samples of pure spruce and mixed spruce-oak stands in southern Sweden as mycelia grown into sand-filled mesh bags placed at three different soil depths (0–10, 10–20 and 20–30 cm). The mesh bags were collected after 12 months and we found that 590±70 kg ha^–1 year^–1 of pure mycelia was produced in spruce stands and 420±160 kg ha^–1 year^–1 in mixed stands. The production of EM mycelia in the mesh bags decreased with soil depth in both stand types but tended to be more concentrated in the top soil in the mixed stands compared to the spruce stands. The fungal biomass was also determined in soil samples taken from different depths by using phospholipid fatty acids as markers for fungal biomass. Subsamples were incubated at 20°C for 5 months and the amount of fungal biomass that degraded during the incubation period was used as an estimate of EM fungal biomass. The EM biomass in the soil profile decreased with soil depth and did not differ significantly between the two stand types. The total EM biomass in the pure spruce stands was estimated to be 4.8±0.9×10³ kg ha^–1 and in the mixed stands 5.8±1.1×10³ kg ha^–1 down to 70 cm depth. The biomass and production estimates of EM mycelia suggest a very long turnover time or that necromass has been included in the biomass estimates. The amount of N present in EM mycelia was estimated to be 121 kg N ha^–1 in spruce stands and 187 kg N ha^–1 in mixed stands. The ¹³C value for mycelia in mesh bags was not influenced by soil depth, indicating that the fungi obtained all their carbon from the tree roots. The ¹³C values in mycelia collected from mixed stands were intermediate to values from pure spruce and pure oak stands suggesting that the EM mycelia received carbon from both spruce and oak trees in the mixed stands. The ¹⁵N value for the EM mycelia and the surrounding soil increased with soil depth suggesting that they obtained their entire N from the surrounding soil.