Effects of Succession on Nitrogen Export in the West-Central Cascades, Oregon

This study examined impacts of succession on N export from 20 headwater stream systems in the west central Cascades of Oregon, a region of low anthropogenic N inputs. The seasonal and successional patterns of nitrate (NO₃−N) concentrations drove differences in total dissolved N concentrations because ammonium (NH₄−N) concentrations were very low (usually < 0.005 mg L⁻¹) and mean dissolved organic nitrogen (DON) concentrations were less variable than nitrate concentrations. In contrast to studies suggesting that DON levels strongly dominate in pristine watersheds, DON accounted for 24, 52, and 51% of the overall mean TDN concentration of our young (defined as predominantly in stand initiation and stem exclusion phases), middle-aged (defined as mixes of mostly understory reinitiation and older phases) and old-growth watersheds, respectively. Although other studies of cutting in unpolluted forests have suggested a harvest effect lasting 5 years or less, our young successional watersheds that were all older than 10 years still lost significantly more N, primarily as NO₃−N, than did watersheds containing more mature forests, even though all forest floor and mineral soil C:N ratios were well above levels reported in the literature for leaching of dissolved inorganic nitrogen. The influence of alder may contribute to these patterns, although hardwood cover was quite low in all watersheds; it is possible that in forested ecosystems with very low anthropogenic N inputs, even very low alder cover in riparian zones can cause elevated N exports. Only the youngest watersheds, with the highest nitrate losses, exhibited seasonal patterns of increased summer uptake by vegetation as well as flushing at the onset of fall freshets. Older watersheds with lower N losses did not exhibit seasonal patterns for any N species. The results, taken together, suggest a role for both vegetation and hydrology in N retention and loss, and add to our understanding of N cycling by successional forest ecosystems influenced by disturbance at various spatial and temporal scales in a region of relatively low anthropogenic N input.     

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.     

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.     

The Long-term Effects of Disturbance on Organic and Inorganic Nitrogen Export in the White Mountains, New Hampshire

Traditional biogeochemical theories suggest that ecosystem nitrogen retention is controlled by biotic N limitation, that stream N losses should increase with successional age, and that increasing N deposition will accelerate this process. These theories ignore the role of dissolved organic nitrogen (DON) as a mechanism of N loss. We examined patterns of organic and inorganic N export from sets of old-growth and historically (80–110 years ago) logged and burned watersheds in the northeastern US, a region of moderate, elevated N deposition. Stream nitrate concentrations were strongly seasonal, and mean (± SD) nitrate export from old-growth watersheds (1.4 ± 0.6 kg N ha⁻¹ y⁻¹) was four times greater than from disturbed watersheds (0.3 ± 0.3 kg N ha⁻¹ y⁻¹), suggesting that biotic control over nitrate loss can persist for a century. DON loss averaged 0.7 (± 0.2) kg N ha⁻¹ y⁻¹ and accounted for 28–87% of total dissolved N (TDN) export. DON concentrations did not vary seasonally or with successional status, but correlated with dissolved organic carbon (DOC), which varied inversely with hardwood forest cover. The patterns of DON loss did not follow expected differences in biotic N demand but instead were consistent with expected differences in DOC production and sorption. Despite decades of moderate N deposition, TDN export was low, and even old-growth forests retained at least 65% of N inputs. The reasons for this high N retention are unclear: if due to a large capacity for N storage or biological removal, N saturation may require several decades to occur; if due to interannual climate variability, large losses of nitrate may occur much sooner. Received 27 April 1999; accepted 30 May 2000.     

Throughfall Nitrogen Deposition Has Different Impacts on Soil Solution Nitrate Concentration in European Coniferous and Deciduous Forests

Increases in the deposition of atmospheric nitrogen (N) influence N cycling in forest ecosystems and can result in negative consequences due to the leaching of nitrate into groundwaters. From December 1995 to February 1998, the Pan-European Programme for the Intensive and Continuous Monitoring of Forest Ecosystems measured forest conditions at a plot scale for conifer and broadleaf forests, including the performance of time series of soil solution chemistry. The influence of various ecosystem conditions on soil solution nitrate concentrations at these forest plots (n = 104) was then analyzed with a statistical model. Soil solution nitrate concentrations varied by season, and summer concentrations were approximately 25% higher than winter ones. Soil solution nitrate concentrations increased dramatically with throughfall (and bulk precipitation) N input for both broadleaf and conifer forests. However, at elevated levels of throughfall N input (more than 10 kg N ha^–1 y^–1), nitrate concentrations were higher in broadleaf than coniferous stands. This tree-specific difference was not observed in response to increased bulk precipitation N input. In coniferous stands, throughfall N input, foliage N concentration, organic layer carbon–nitrogen (C:N) ratio, and nitrate concentrations covaried. Soil solution nitrate concentrations in conifer plots were best explained by a model with throughfall N and organic layer C:N as main factors, where C:N ratio could be replaced by foliage N. The organic layer C:N ratio classes of more than 30, 25–30, and less than 25, as well as the foliage N (mg N g^–1) classes of less than 13, 13–17, and more than 17, indicated low, intermediate, and high risks of nitrate leaching, respectively. In broadleaf forests, correlations between N characteristics were less pronounced, and soil solution nitrate concentrations were best explained by throughfall N and soil pH (0–10-cm depth). These results indicate that the responses of soil solution nitrate concentration to changes in N input are more pronounced in broadleaf than in coniferous forests, because in European forests broadleaf species grow on the more fertile soils.     

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.     

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.     

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.     

The influence of watershed characteristics on nitrogen export to and marine fate in Hood Canal,Washington, USA

Hood Canal, Washington, USA, is a poorly ventilated fjord-like sub-basin of Puget Sound that commonly experiences hypoxia. This study examined the influence of watershed soils, vegetation, physical features, and population density on nitrogen (N) export to Hood Canal from 43 tributaries. We also linked our watershed study to the estuary using a salinity mass balance model that calculated the relative magnitude of N loading to Hood Canal from watershed, direct precipitation, and marine sources. The overall flow-weighted total dissolved N (TDN) and particulate N input concentrations to Hood Canal were 152 and 49 μg l⁻¹, respectively. Nitrate and dissolved organic N comprised 64 and 29% of TDN, respectively. The optimal regression models for TDN concentration and areal yield included a land cover term suggesting an effect of N-fixing red alder (Alnus rubra) and a human population density term (suggesting onsite septic system (OSS) discharges). There was pronounced seasonality in stream water TDN concentrations, particularly for catchments with a high prevalence of red alder, with the lowest concentrations occurring in the summer and the highest occurring in November–December. Due to strong seasonality in TDN concentrations and in particular stream flow, over 60% of the TDN export from this watershed occurred during the 3 month period of November–January. Entrainment of marine water into the surface layer of Hood Canal accounted for ≈98% of N loading to the euphotic zone of this estuary, and in a worst case scenario OSS N inputs contribute ≈0.5% of total N loading. Domestic wastewater discharges and red alders appear to be a very important N source for many streams, but a minor nutrient source for the estuary as a whole.     

Cation distribution,cycling, and removal from mineral soil in Douglas-fir and red alder forests

Overstory species influence the distribution and dynamics of nutrients in forest ecosystems. Ecosystem-level estimates of Ca, Mg, and K pools and cycles in 50-year old Douglas-fir and red alder stands were used to determine the effect of overstory composition on net cation removal from the mineral soil, i.e. cation export from the soil in excess of additions. Net cation removal from Douglas-fir soil was 8 kg Ca ha^–1 yr^–1, 1 kg Mg ha^–1 yr^–1, and 0.3 kg K ha^–1 yr^–1. Annual cation export from soil by uptake and accumulation in live woody tissue and O horizon was of similar magnitude to leaching in soil solution. Atmospheric deposition partially off-set export by adding cations equivalent to 28–88% of cation export. Net cation removal from red alder soil was 58 kg Ca ha^–1 yr^–1, 9 kg Mg ha^–1 yr^–1, and 11 kg K ha^–1 yr^–1. Annual cation accumulation in live woody tissue and O horizon was three times greater than in Douglas-fir, while cation leaching in soil solution was five to eight times greater. The lack of excessive depletion of exchangeable cations in the red alder soil suggests that mineral weathering, rather than exchangeable cations, was the source of most of the removed cations. Nitric acid generated during nitrification in red alder soil led to high rates of weathering and NO₃-driven cation leaching.     

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.     

Nitrogen input–output budgets for lake-containing watersheds in the Adirondack region of New York

The Adirondack region of New York is characterized by soils and surface waters that are sensitive to inputs of strong acids, receiving among the highest rates of atmospheric nitrogen (N) deposition in the United States. Atmospheric N deposition to Adirondack ecosystems may contribute to the acidification of soils through losses of exchangeable basic cations and the acidification of surface waters in part due to increased mobility of nitrate (NO₃^–). This response is particularly evident in watersheds that exhibit nitrogen saturation. To evaluate the contribution of atmospheric N deposition to the N export and the capacity of lake-containing watersheds to remove, store, or release N, annual N input–output budgets were estimated for 52 lake-containing watersheds in the Adirondack region from 1998 to 2000. Wet N deposition was used as the N input and the lake N discharge loss was used as the N output based on modeled hydrology and measured monthly solute concentrations. Annual outputs were also estimated for dissolved organic carbon (DOC). Wet N deposition increased from the northeast to the southwest across the region. Lake N drainage losses, which exhibited a wider range of values than wet N deposition, did not show any distinctive spatial pattern, although there was some evidence of a relationship between wet N deposition and the lake N drainage loss. Wet N deposition was also related to the fraction of N removed or retained within the watersheds (i.e., the fraction of net N hydrologic flux relative to wet N deposition, calculated as [(wet N deposition minus lake N drainage loss)/wet N deposition]). In addition to wet N deposition, watershed attributes also had effects on the exports of NO₃^–, ammonium (NH₄⁺), dissolved organic nitrogen (DON), and DOC, the DOC/DON export ratio, and the N flux removed or retained within the watersheds (i.e., net N hydrologic flux, calculated as [wet N deposition less lake N drainage loss]). Elevation was strongly related with the lake drainage losses of NO₃^–, NH₄⁺, and DON, net NO₃^– hydrologic flux (i.e., NO₃^– deposition less NO₃^– drainage loss), and the fraction of net NO₃^– hydrologic flux, but not with the DOC drainage loss. Both DON and DOC drainage losses from the lakes increased with the proportion of watershed area occupied by wetlands, with a stronger relationship for DOC. The effects of wetlands and forest type on NO₃^– flux were evident for the estimated NO₃^– fluxes flowing from the watershed drainage area into the lakes, but were masked in the drainage losses flowing out of the lakes. The DOC/DON export ratios from the lake-containing watersheds were in general lower than those from forest floor leachates or streams in New England and were intermediate between the values of autochthonous and allochthonous dissolved organic matter (DOM) reported for various lakes. The DOC/DON ratios for seepage lakes were lower than those for drainage lakes. In-lake processes regulating N exports may include denitrification, planktonic depletion, degradation of DOM, and the contribution of autochthonous DOM and the influences of in-lake processes were also reflected in the relationships with hydraulic retention time. The N fluxes removed or stored within the lakes substantially varied among the lakes. Our analysis demonstrates that for these northern temperate lake-containing watershed ecosystems, many factors, including atmospheric N deposition, landscape features, hydrologic flowpaths, and retention in ponded waters, regulated the spatial patterns of net N hydrologic flux within the lake-containing watersheds and the loss of N solutes through drainage waters.     

Biotic versus hydrologic control over seasonal nitrate leaching in a floodplain forest

Strong seasonal increases in aquatic (stream, ground and hyporheic water) nitrate have been observed in a variety of ecosystems. In most cases, changes in hydrological and vegetative activity occur contemporaneously, making it difficult to determine whether soil leaching is being driven by increases in the availability of leachable N or is simply due to flushing of N that has accumulated over longer periods. Three studies were conducted to better determine controls on soil nitrate leaching in a near-pristine temperate floodplain ecosystem receiving large N inputs via N-fixation by red alder: 1) an artificial rainfall experiment was conducted to estimate N-leaching potential during the summer, when plant uptake is high and new inputs of organic matter are low; 2) soil solution, groundwater and surface water were sampled during a major autumn storm to document exchanges at the seasonal transition, when plant uptake is low and inputs of senescent organic matter are high; and 3) monthly samples of soil and aquatic nitrogen were collected in 1997 and 1998 to document seasonal patterns of N exchanges. Collectively, these studies demonstrate the importance of hydrologic factors in controlling N flux. Nitrate was rapidly leached from soils during actual and simulated rainstorms. Two pathways of nitrate leaching were identified. Localized flooding and direct leaching of streamside soils into surface waters contributed to high solute concentrations in peak flows. Nitrate that leached into interstitial waters was subject to various factors that could delay or reduce its delivery to surface waters. Greater residence time may increase the influence of this component of stormflow on ecosystem productivity. While soil nitrate pools were rapidly depleted during rainstorms, accumulation of soil nitrate occurred over summer dry periods. Large differences in soil and aquatic nitrate concentrations between two years with contrasting rainfall highlight the potential for inter-annual hydrologic variability to affect ecosystem nutrient cycling.     

Nitrate leaching from three afforestation chronosequences on former arable land in Denmark

In regions dominated by agricultural activities, nitrogen (N) is recognized as a major pollutant in aquatic environments. In north‐western Europe, afforestation of agricultural land is part of a strategy to improve water quality. In Denmark, former arable land has been afforested during the past 40–50 years. This study evaluated the effect of afforestation of former arable land on nitrate leaching, based on three afforestation chronosequences. Precipitation, canopy throughfall and soil water were collected and soil moisture was monitored at two Danish locations, Vestskoven (nutrient‐rich, medium deposition) and Gejlvang (nutrient‐poor, high deposition). Afforestation was performed using Norway spruce [Picea abies (Karst.) L.] and common oak (Quercus robur L.) at Vestskoven and Norway spruce at Gejlvang. The results suggest that afforestation of former arable land initially leads to lower nitrate leaching than that occurring under the former agricultural land use, and largely below the standard of 50 mg NO⁻₃ L⁻¹ for groundwater to be utilized as drinking water. Nitrate concentrations became almost negligible in forest stands of 5–20 years of age. However, after canopy closure (>20 years) nitrate concentrations below the root zone and nitrate leaching tended to increase. This was attributed to increased N deposition with increasing canopy development and decreased N demand once the most N‐rich biomass compartments had been built up. Nitrate leaching started to increase at a throughfall deposition level of about 10 kg N ha⁻¹ yr⁻¹. Compared with nutrient‐poor sandy soils, nutrient‐rich clayey soils appeared more vulnerable to disturbance of the N cycle and to increased N deposition, leading to N saturation and enhanced nitrate leaching. In approximately the first 35 years after afforestation, nitrate leaching below the root zone was generally higher below oak than below Norway spruce.     

Earthworms Increase Nitrogen Leaching to Greater Soil Depths in Row Crop Agroecosystems

Many biological functions of soil organisms are replaced in intensive agricultural systems, but earthworms and other soil invertebrates may continue to have significant effects on nutrient cycling in these disturbed systems. We investigated the influence of earthworms on leaching of water and nitrogen in corn (Zea mays L.) agroecosystems in a long-term (6-year) field experiment in Wooster, Ohio, USA. We employed a split-plot experimental design in which main plots received one of three nutrient treatments (cow manure, legume–grass mixture, inorganic fertilizer) and contained three 4.5 × 4.5-m field enclosures in which earthworm populations were increased, decreased, or unmodified. We installed zero-tension lysimeters beneath enclosures with increased or decreased populations and collected leachates regularly in 1996, analyzing them for water volume and concentrations of NH₄⁺, NO₃^–, and dissolved organic nitrogen (DON). Earthworms did not influence concentrations of inorganic N or DON but greatly increased leachate volume. The total flux of N in soil leachates was 2.5-fold greater in plots with increased earthworm populations than in those with decreased populations. Earthworm population density was positively correlated with total N leaching flux (r² = 0.49). Leaching losses of N to a depth of 45 cm were greater in the inorganically fertilized than in the organically fertilized plots, possibly due to greater inorganic N concentrations and lower immobilization potential in inorganically fertilized systems. Our results indicate that earthworms can increase the leaching of water and nitrogen to greater soil depths, potentially increasing N leaching from the system.Present address: Departamento de Ecoloxia e Bioloxía Animal, Universidade de Vigo, E-36200, Spain. Present address: Archbold Biological Station, 300 Buck Island Ranch Rd., Lake Placid, Florida 33852, USA. ¶Present address: P.O. Box 303, Yucca Valley, California 92286, USA.     

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.     

Differences among maize cultivars in the utilization of soil nitrate and the related losses of nitrate through leaching

In a 2-year field experiment conducted on a Gleyic Luvisol in Stuttgart-Hohenheim one experimental and nine commercial maize cultivars were compared for their ability to utilize soil nitrate and to reduce related losses of nitrate through leaching. Soil nitrate was monitored periodically in CaCl₂ extracts and in suction cup water. Nitrate concentrations in suction water were generally higher than in CaCl₂ extracts. Both methods revealed that all cultivars examined were able to extract nitrate down to a soil depth of at least 120 cm (1988 season) or 150 cm (1987 season). Significant differences among the cultivars existed in nitrate depletion particularly in the subsoil. At harvest, residual nitrate in the upper 150 cm of the profile ranged from 73–110 kg N ha^–1 in 1987 and from 59–119 kg N ha^–1 in 1988. Residual nitrate was closely correlated with nitrate losses by leaching because water infiltration at 120 cm soil depth started 4 weeks after harvest (1987) or immediately after harvest (1988) and continued until early summer of the following year. The calculated amount of nitrate lost by leaching was strongly influenced by the method of calculation. During the winter of 1987/88 nitrate leaching ranged from 57–84 kg N ha^–1 (suction cups) and 40–55 kg N ha^–1 (CaCl₂ extracts), respectively. The corresponding values for the winter of 1988/89 were 47–79 and 20–39 kg N ha^–1, respectively. ei]Section editor: B E Clothier     

Nitrogen Dynamics in Ice Storm-Damaged Forest Ecosystems: Implications for Nitrogen Limitation Theory

Despite the widely recognized importance of disturbance in accelerating the loss of elements from land, there have been few empirical studies of the effects of natural disturbances on nitrogen (N) dynamics in forest ecosystems. We were provided the unusual opportunity for such study, partly because the intensively monitored watersheds at the Hubbard Brook Experimental Forest (HBEF), New Hampshire, experienced severe canopy damage following an ice storm. Here we report the effects of this disturbance on internal N cycling and loss for watershed 1 (W1) and watershed 6 (W6) at the HBEF and patterns of N loss from nine other severely damaged watersheds across the southern White Mountains. This approach allowed us to test one component of N limitation theory, which suggests that N losses accompanying natural disturbances can lead to the maintenance of N limitation in temperate zone forest ecosystems. Prior to the ice storm, fluxes of nitrate (NO₃ ^–) at the base of W1 and W6 were similar and were much lower than N inputs in atmospheric deposition. Following the ice storm, drainage water NO₃ ^– concentrations increased to levels that were seven to ten times greater than predisturbance values. We observed no significant differences in N mineralization, nitrification, or denitrification between damaged and undamaged areas in the HBEF watersheds, however. This result suggests that elevated NO₃ ^- concentrations were not necessarily due to accelerated rates of N cycling by soil microbes but likely resulted from decreased plant uptake of NO₃ ^-. At the regional scale, we observed high variability in the magnitude of NO₃ ^- losses: while six of the surveyed watersheds showed accelerated rates of NO₃ ^– loss, three did not. Moreover, in contrast to the strong linear relationship between NO₃ ^– loss and crown damage within HBEF watersheds [r ²: (W1 = 0.91, W6 = 0.85)], stream water NO₃ ^– concentrations were weakly related to crown damage (r ² = 0.17) across our regional sites. The efflux of NO₃ ^– associated with the ice storm was slightly higher than values reported for soil freezing and insect defoliation episodes, but was approximately two to ten times lower than NO₃ ^– fluxes associated with forest harvesting. Because over one half of the entire years worth of N deposition was lost following the ice storm, we conclude that catastrophic disturbances contribute synergistically to the maintenance of N limitation and widely observed delays of N saturation in northern, temperate zone forest ecosystems. Present address: Department of Ecology and Evolutionary Biology, Princeton University, Guyot Hall, Princeton, New Jersey 08544, USA.     

Effects of soil freezing disturbance on soil solution nitrogen, phosphorus, and carbon chemistry in a northern hardwood ecosystem

Reductions in snow cover undera warmer climate may cause soil freezing eventsto become more common in northern temperateecosystems. In this experiment, snow cover wasmanipulated to simulate the late development ofsnowpack and to induce soil freezing. Thismanipulation was used to examine the effects ofsoil freezing disturbance on soil solutionnitrogen (N), phosphorus (P), and carbon (C)chemistry in four experimental stands (twosugar maple and two yellow birch) at theHubbard Brook Experimental Forest (HBEF) in theWhite Mountains of New Hampshire. Soilfreezing enhanced soil solution Nconcentrations and transport from the forestfloor. Nitrate (NO₃ ^–) was thedominant N species mobilized in the forestfloor of sugar maple stands after soilfreezing, while ammonium (NH₄ ⁺) anddissolved organic nitrogen (DON) were thedominant forms of N leaching from the forestfloor of treated yellow birch stands. Rates ofN leaching at stands subjected to soil freezingranged from 490 to 4,600 mol ha^–1yr^–1, significant in comparison to wet Ndeposition (530 mol ha^–1 yr^–1) andstream NO₃ ^– export (25 mol ha^–1yr^–1) in this northern forest ecosystem. Soil solution fluxes of P_i from the forestfloor of sugar maple stands after soil freezingranged from 15 to 32 mol ha^–1 yr^–1;this elevated mobilization of P_i coincidedwith heightened NO₃ ^– leaching. Elevated leaching of P_i from the forestfloor was coupled with enhanced retention ofP_i in the mineral soil Bs horizon. Thequantities of P_i mobilized from the forestfloor were significant relative to theavailable P pool (22 mol ha^–1) as well asnet P mineralization rates in the forest floor(180 mol ha^–1 yr^–1). Increased fineroot mortality was likely an important sourceof mobile N and P_i from the forest floor,but other factors (decreased N and P uptake byroots and increased physical disruption of soilaggregates) may also have contributed to theenhanced leaching of nutrients. Microbialmortality did not contribute to the acceleratedN and P leaching after soil freezing. Resultssuggest that soil freezing events may increaserates of N and P loss, with potential effectson soil N and P availability, ecosystemproductivity, as well as surface wateracidification and eutrophication.     

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.