Nitrogen mineralization,nitrification and denitrification in upland and wetland ecosystems

Summary Nitrogen mineralization, nitrification, denitrification, and microbial biomass were evaluated in four representative ecosystems in east-central Minnesota. The study ecosystems included: old field, swamp forest, savanna, and upland pin oak forest. Due to a high regional water table and permeable soils, the upland and wetland ecosystems were separated by relatively short distances (2 to 5 m). Two randomly selected sites within each ecosystem were sampled for an entire growing season. Soil samples were collected at 5-week intervals to determine rates of N cycling processes and changes in microbial biomass. Mean daily N mineralization rates during five-week in situ soil incubations were significantly different among sampling dates and ecosystems. The highest annual rates were measured in the upland pin oak ecosystem (8.6 g N m^–2 yr^–1), and the lowest rates in the swamp forest (1.5 g N m^–2 yr^–1); nitrification followed an identical pattern. Denitrification was relatively high in the swamp forest during early spring (8040 g N₂O–N m^–2 d^–1) and late autumn (2525 g N₂O–N m^–2 d^–1); nitrification occurred at rates sufficient to sustain these losses. In the well-drained uplands, rates of denitrification were generally lower and equivalent to rates of atmospheric N inputs. Microbial C and N were consistently higher in the swamp forest than in the other ecosystems; both were positively correlated with average daily rates of N mineralization. In the subtle landscape of east-central Minnesota, rates of N cycling can differ by an order of magnitude across relatively short distances.     

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.     

The effect of the presence of a forage legume on nitrogen and carbon levels in soils under Brachiaria pastures in the Atlantic forest region of the South of Bahia,Brazil

The impact of forest clearance, and its replacement by Brachiaria pastures, on soil carbon reserves has been studied at many sites in the Brazilian Amazonia, but to date there appear to be no reports of similar studies undertaken in the Atlantic forest region of Brazil. In this study performed in the extreme south of Bahia, the changes in C and N content of the soil were evaluated from the time of establishment of grass-only B. humidicola and mixed B. humidicola/Desmodium ovalifolium pastures through 9 years of grazing in comparison with the C and N contents of the adjacent secondary forest. The decline in the content of soil C derived from the forest (C3) vegetation and the accumulation of that derived from the Brachiaria (C4) were followed by determining the ¹³C natural abundance of the soil organic matter (SOM). The pastures were established in 1987, 10 years after deforestation, and it was estimated that until 1994 there was a loss in forest-derived C in the top 30 cm of soil of approximately 20% (9.1 Mg C ha^–1). After the establishment of the pastures, C derived from Brachiaria accumulated steadily such that at the final sampling (1997) it was estimated 13.9 Mg ha^–1 was derived from this source under the grass-only pasture (0–30 cm). Samples taken from all pastures and the forest in 1997 to a depth of 100 cm showed that below 40 cm depth there was no significant contribution of the Brachiaria-derived C and that total C reserves under the grass/legume and the grass-only pastures were slightly higher than under the forest (not significant at P=0.05). The more detailed sampling under the pastures showed that to a depth of 30 cm there was significantly (P<0.05) more C under the mixed pasture than the grass-only pasture. It was estimated that from the time of establishment the apparent rate of C accumulation (0–100 cm depth) under the grass/legume pastures (1.17 Mg ha^–1 yr^–1) was almost double that under the grass-only pastures (0.66 Mg ha^–1 yr^–1). The data indicated that newly incorporated SOM derived from the Brachiaria had a considerably higher C:N ratio than that present under the forest.     

The fate of15N enriched throughfall in two coniferous forest stands at different nitrogen deposition levels

The stable isotope¹⁵N was added as (¹⁵NH₄)₂SO₄ to throughfall water for one year, to study the fate of the deposited nitrogen at different levels of N deposition in two N saturated coniferous forests ecosystems in the Netherlands. The fate of the¹⁵N was followed at high-N (44–55 kg N ha^–1 yr^–1) 1) and low-N (4–6 kg N ha^–1 yr^–1) deposition in plots established under transparent roofs build under the canopy in a Douglas fir (Pseudotsuga menziesii (Mirb.) Franco.) and Scots pine (Pinus sylvestris L.) forest.The applied¹⁵N was detectable in needles and twigs, the soil and soil water leaching below the rooting zone (90 cm depth). Total¹⁵N recovery in major ecosystem compartments was 71–100% during two successive growing seasons after the start of a year-round¹⁵N application to throughfall-N. Nine months after the year-round¹⁵N application, the¹⁵N assimilated into tree biomass was 29–33% of the¹⁵N added in the Douglas fir stand and less than 17% in the Scots pine stand. At the same time total¹⁵N retention in the soil (down to 70 cm) of the high-N plots was about 37% of the deposited¹⁵NH₄-N, whereas 46% and 65% of the¹⁵N was found in the soil of the low-N deposition plots at the Douglas fir and Scots pine stand, respectively. The organic layers accounted for 60% of the¹⁵N retained in the soil. The total N deposition exceeded the demand of the vegetation and microbial immobilization. Total¹⁵N leaching losses within a year (below 90 cm) were 10–20% in the high-N deposition plots in comparison to 2–6% in the lowered nitrogen input plots. Relative retention in the soil and vegetation increased at lower N-input levels.Species differences in uptake and tree health seem to contribute to lower¹⁵N recoveries in the Scots pine trees compared to the Douglas fir trees. The excessive N deposition and resulting N saturation lead to conditions were the health and functioning of biota were negatively influenced. At decreased N deposition, lower leaching losses together with increased soil and plant retention indicated a change in the fate of the¹⁵N deposited. This may have resulted from changes in ecosystem processes, and thus a shift along the continuum of N saturation to N limitation.     

Nitrogen cycling in a northern hardwood forest: Do species matter?

To investigate the influence of individual tree species on nitrogen (N) cycling in forests, we measured key characteristics of the N cycle in small single-species plots of five dominant tree species in the Catskill Mountains of New York State. The species studied were sugar maple (Acer saccharum), American beech (Fagus grandifolia), yellow birch (Betula alleghaniensis), eastern hemlock (Tsuga canadensis), and red oak (Quercus rubra). The five species varied markedly in N cycling characteristics. For example, hemlock plots consistently showed characteristics associated with "slow" N cycling, including low foliar and litter N, high soil C:N, low extractable N pools, low rates of potential net N mineralization and nitrification and low NO ₃ ^– amounts trapped in ion-exchange resin bags buried in the mineral soil. Sugar maple plots had the lowest soil C:N, and the highest levels of soil characteristics associated with NO ₃ ^– production and loss (nitrification, extractable NO ₃ ^– , and resin bag NO ₃ ^– ). In contrast, red oak plots had near-average net mineralization rates and soil C:N ratios, but very low values of the variables associated with NO ₃ ^– production and loss. Correlations between soil N transformations and litter concentrations of N, lignin, lignin:N ratio, or phenolic constituents were generally weak. The inverse correlation between net nitrification rate and soil C:N that has been reported in the literature was present in this data set only if red oak plots were excluded from the analysis. This study indicates that tree species can exert a strong control on N cycling in forest ecosystems that appears to be mediated through the quality of soil organic matter, but that standard measures of litter quality cannot explain the mechanism of control.     

Soil N mineralization and nitrification in relation to nitrogen solution chemistry in a small forested watershed

Spatial variations in soil processes regulating mineral N losses to streams were studied in a small watershed near Toronto, Ontario. Annual net N mineralization in the 0–8 cm soil was measured in adjacent upland and riparian forest stands using in situ soil incubations from April 1985 to 1987. Mean annual rates of soil N mineralization and nitrification were higher in a maple soil (93.8 and 87.0 kg.ha^–1) than in a pine soil (23.3 and 8.2 kg.ha^–1 ). Very low mean rates of mineralization (3.3 kg.ha^–1) and nitrification (3.4 kg.ha^–1) were found in a riparian hemlock stand. Average NO₃-N concentrations in soil solutions were 0.3–1.0 mg.L^–1 in the maple stand and >0.06mg.L^–1 in the pine stand. Concentrations of NO₃–N in shallow ground water and stream water were 3–4× greater in a maple subwatershed than in a pine subwatershed. Rapid N uptake by vegetation was an important mechanism reducing solution losses of NO₃–N in the maple stand. Low rates of nitrification were mainly responsible for negligible NO₃–N solution losses in the pine stand.     

Natural<Superscript>15</Superscript> N Abundance of Plants and Soil N in a Temperate Coniferous Forest

Measurement of nitrogen isotopic composition (¹⁵N) of plants and soil nitrogen might allow the characteristics of N transformation in an ecosystem to be detected. We tested the measurement of ¹⁵N for its ability to provide a picture of N dynamics at the ecosystem level by doing a simple comparison of ¹⁵N between soil N pools and plants, and by using an existing model. ¹⁵N of plants and soil N was measured together with foliar nitrate reductase activity (NRA) and the foliar NO₃^– pool at two sites with different nitrification rates in a temperature forest in Japan. ¹⁵N of plants was similar to that of soil NO₃^– in the high-nitrification site. Because of high foliar NRA and the large foliar NO₃^– pool at this site, we concluded that plant ¹⁵N indicated a great reliance of plants on soil NO₃^– there. However, many ¹⁵N of soil N overlapped each other at the other site, and ¹⁵N could not provide definitive evidence of the N source. The existing model was verified by measured ¹⁵N of soil inorganic N and it explained the variations of plant ¹⁵N between the two sites in the context of relative importance of nitrification, but more information about isotopic fractionations during plant N uptake is required for quantitative discussions about the plant N source. The model applied here can provide a basis to compare ¹⁵N signatures from different ecosystems and to understand N dynamics.     

The nitrogen cycle in lodgepole pine forests,southeastern Wyoming

Storage and flux of nitrogen were studied in several contrasting lodgepole pine (Pinus contorta spp.latifolia) forests in southeastern Wyoming. The mineral soil contained most of the N in these ecosystems (range of 315–860 g · m^–2), with aboveground detritus (37.5–48.8g · m^–2) and living biomass (19.5–24.0 g · m^–2) storing much smaller amounts. About 60–70% of the total N in vegetation was aboveground, and N concentrations in plant tissues were unusually low (foliage = 0.7% N), as were N input via wet precipitation (0.25 g · m^–2 · yr^–1), and biological fixation of atmospheric N (<0.03 g · m^–2 · yr^–1, except locally in some stands at low elevations where symbiotic fixation by the leguminous herbLupinus argenteus probably exceeded 0.1 g · m^–2 · yr^–1).Because of low concentrations in litterfall and limited opportunity for leaching, N accumulated in decaying leaves for 6–7 yr following leaf fall. This process represented an annual flux of about 0.5g · m^–2 to the 01 horizon. Only 20% of this flux was provided by throughfall, with the remaining 0.4g · m^–2 · yr^–1 apparently added from layers below. Low mineralization and small amounts of N uptake from the 02 are likely because of minimal rooting in the forest floor (as defined herein) and negligible mineral N (< 0.05 mg · L^–1) in 02 leachate. A critical transport process was solubilization of organic N, mostly fulvic acids. Most of the organic N from the forest floor was retained within the major tree rooting zone (0–40 cm), and mineralization of soil organic N provided NH₄ for tree uptake. Nitrate was at trace levels in soil solutions, and a long lag in nitrification was always observed under disturbed conditions. Total root nitrogen uptake was calculated to be 1.25 gN · m^–2 · yr^–1 with estimated root turnover of 0.37-gN · m^–2 · yr^–1, and the soil horizons appeared to be nearly in balance with respect to N. The high demand for mineralized N and the precipitation of fulvic acid in the mineral soil resulted in minimal deep leaching in most stands (< 0.02 g · m^–2 · yr^–1). These forests provide an extreme example of nitrogen behavior in dry, infertile forests.     

Phosphorus composition of upland soils polluted by long-term atmospheric nitrogen deposition

Atmospheric N deposition can enhance biological P limitation in terrestrial ecosystems and increase the importance of organic P to plants and microorganisms. We used NaOH–EDTA extraction and solution ³¹P NMR spectroscopy to determine the P composition of soils in the Upper Teesdale National Nature Reserve, northern England, an upland region influenced by such deposition for at least 150 years. Three characteristic soil types were sampled on three occasions during an annual cycle: blanket peat (318 mg g^–1 total C, 607 g g^–1 total P, pH 3.9); acid organic soil under grassland (354 mg g^–1 total C, 1190 g g^–1 total P, pH 3.7); calcareous soil under grassland (140 mg g^–1 total C, 649 g g^–1 total P, pH 7.3). Between 58 and 99% of the total P in soil and litter layers was extracted by 0.25 M NaOH + 0.05 M EDTA. Extracts of all soils were dominated by organic P, mainly in the form of orthophosphate monoesters (43–69% extracted P). The two acidic soils also contained large proportions of orthophosphate diesters (6–19% extracted P) and phosphonates (7–16% extracted P), suggesting that these compounds become stabilised at low pH. However, a seasonal trend of increasing orthophosphate monoester-to-diester ratios, most evident in the calcareous grassland soil, indicated the preferential degradation of orthophosphate diesters during the growing season. Orthophosphate was the major inorganic P compound (17–34% extracted P), and all soils contained pyrophosphate (1–5% extracted P). However, orthophosphate determined in the NaOH–EDTA extracts by solution ³¹P NMR spectroscopy was substantially greater than that determined by molybdate colourimetry, suggesting that orthophosphate occurred in complexes with humic compounds that were not detected by conventional procedures. Our results suggest that organisms able to use recalcitrant soil organic P may have a competitive advantage in environments under enhanced atmospheric N deposition.     

Vertical transport of dissolved organic C and N under long-term N amendments in pine and hardwood forests

At the Harvard Forest, Massachusetts, a long-term effort is under way to study responses in ecosystem biogeochemistry to chronic inputs of N in atmospheric deposition in the region. Since 1988, experimental additions of NH₄NO₃ (0, 5 and 15 g N m^–2 yr^–1) have been made in two forest stands:Pinus resinosa (red pine) and mixed hardwood. In the seventh year of the study, we measured solute concentrations and estimated solute fluxes in throughfall and at two soil depths, beneath the forest floors (Oa) and beneath the B horizons.Beneath the Oa, concentrations and fluxes of dissolved organic C and N (DOC and DON) were higher in the coniferous stand than in the hardwood stand. The mineral soil exerted a strong homogenizing effect on concentrations beneath the B horizons. In reference plots (no N additions), DON composed 56% (pine) and 67% (hardwood) of the total dissolved nitrogen (TDN) transported downward from the forest floor to the mineral soil, and 98% of the TDN exported from the solums. Under N amendments, fluxes of DON from the forest floor correlated positively with rates of N addition, but fluxes of inorganic N from the Oa exceeded those of DON. Export of DON from the solums appeared unaffected by 7 years of N amendments, but as in the Oa, DON composed smaller fractions of TDN exports under N amendments. DOC fluxes were not strongly related to N amendment rates, but ratios of DOC:DON often decreased.The hardwood forest floor exhibited a much stronger sink for inorganic N than did the pine forest floor, making the inputs of dissolved N to mineral soil much greater in the pine stand. Under the high-N treatment, exports of inorganic N from the solum of the pine stand were increased >500-fold over reference (5.2 vs. 0.01 g N m^–2 yr^–1), consistent with other manifestations of nitrogen saturation. Exports of N from the solum in the pine forest decreased in the order NO₃-N> NH₄-N> DON, with exports of inorganic N 14-fold higher than exports of DON. In the hardwood forest, in contrast, increased sinks for inorganic N under N amendments resulted in exports of inorganic N that remained lower than DON exports in N-amended plots as well as the reference plot.     

Effects of pasture introduction on soil CO2 emissions during the dry season in the state of Rondônia,Brazil

Soil CO₂ evolution rates, soil temperatures and moisture were measured during the dry season in two forest-to-pasture chronosequences in Rondônia, Brazil. The study included pastures ranging from 3 to 80 years-old. Mean dry-season CO₂ evolution from the forest in chronosequence 1, 88.8 mg CO₂-C m^–2h^–1 was lower than from the pastures which ranged from 111 to 158 mg CO₂-C m^–2h^–1. We found that temperature was not a good predictor of CO₂ emissions from pasture but that there was a significant relationship (r = 0.72,p < 0.05) between soil moisture and pasture emissions. The ¹³C of the soil CO₂ emissions also was measured on chronosequence I; ¹³C of the CO₂ emitted from the C3 forest was –29.43%. Pasture¹³CO₂ values increased from –17.91%. in the 3 year-old pasture to –12.86% in the 80 year-old, reflecting the increasing C₄ inputs with pasture age. Even in the youngest (3 year-old) pasture, 70 percent of the CO₂ evolved originated from C₄ pasture-derived carbon.     

15N abundance of surface soils,roots and mycorrhizas in profiles of European forest soils

¹⁵N natural abundances of soil total N, roots and mycorrhizas were studied in surface soil profiles in coniferous and broadleaved forests along a transect from central to northern Europe. Under conditions of N limitation in Sweden, there was an increase in ¹⁵N of soil total N of up to 9% from the uppermost horizon of the organic mor layer down to the upper 0–5 cm of the mineral soil. The ¹⁵N of roots was only slightly lower than that of soil total N in the upper organic horizon, but further down roots were up to 5% depleted under such conditions. In experimentally N-enriched forest in Sweden, i.e. in plots which have received an average of c. 100 kg N ha^–1 year^–1 for 20 years and which retain less than 50% of this added N in the stand and the soil down to 20 cm depth, and in some forests in central Europe, the increase in ¹⁵N with depth in soil total N was smaller. An increase in ¹⁵N of the surface soil was even observed on experimentally N-enriched plots, although other data suggest that the N fertilizer added was depleted in¹⁵N. In such cases roots could be enriched in¹⁵N relative to soil total N, suggesting that labelling of the surface soil is via the pathway: — available pools of N-plant N-litter N. Under N-limiting conditions roots of different species sampled from the same soil horizon showed similar ¹⁵N. By contrast, in experimentally N-enriched forest ¹⁵N of roots increased in the sequence: ericaceous dwarf shrubs15N enriched compounds in fungal material, which could contribute to explain the observed ¹⁵N profiles if fungal material is enriched, because it is a precursor of stable organic matter and recalcitrant N. This could act in addition to the previous explanation of the isotopically lighter soil surface in forests: plant uptake of ¹⁵N-depleted N and its redeposition onto the soil surface by litter-fall.     

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.     

The role of monoterpenes in soil nitrogen cycling processes in ponderosa pine

The effects of select monoterpenes on nitrogen (N) mineralization and nitrification potentials were determined in four separate laboratory bioassays. The effect of increasing monoterpene addition was an initial reduction in NO₃ ^–-N production (nitrification inhibition), followed by a reduction in the sum of NH₄ ⁺-N and NO₃ ^–-N (inhibition of net N mineralization and net immobilization at high monoterpene additions. Monoterpenes could produce this pattern by inhibiting nitrification, reducing net N mineralization, enhancing immobilization of NO₃ ^–-N relative to NH₄ ⁺-N, and/or stimulating overall net immobilization of N by carbon-rich material.Initial monoterpene concentrations in the assay soils were about 5% of the added amount and were below detection after incubation in most samples.Potential N mineralization-immobilization, nitrification, and soil monoterpene concentrations were determined by soil horizon for four collections from a ponderosa pine (Pinus ponderosa) stand in New Mexico. Concentrations of monoterpenes declined exponentially with soil depth and varied greatly within a horizon. Monoterpene content of the forest floor was not correlated with forest floor biomass. Net N mineralization was inversely correlated with total monoterpene content of all sampled horizons. Nitrification was greatest in the mineral soil, intermediate in the F-H horizon, and never occurred in the L horizon. Nitrification in the mineral soil was inversely correlated with the amount of monoterpenes in the L horizon that contain terminal unsaturated carbon-carbon bonds (r ² = 0.37, P 0.01). This pattern in the field corresponded to the pattern shown in the laboratory assays with increasing monoterpene additions.     

Nitrogen biogeochemistry of three hardwood ecosystems in the Adirondack Region of New York

The biogeochemistry of nitrogen (N)was evaluated for three forest ecosystems[Woods Lake (WL), Pancake-Hall Creek (PHC) andHuntington Forest (HF)] in the Adirondackregion of New York, U.S.A. to evaluate theresponse of a range of N atmospheric inputsand experimental N additions. Bulk Ndeposition was higher at sites in the westthan those in the central and easternAdirondacks. These higher atmospheric N inputswere reflected in higher bulk throughfallfluxes of N (WL and PHC, 10.1 and 12.0 kg Nha^–1 yr^–1, respectively) in thewestern Adirondacks than at HF (4.6 kg Nha^–1 yr^–1) in the centralAdirondacks. Nitrogen was added to plots as(NH₄)₂SO₄ at 14 and 28 kg Nha^–1 yr^–1 or as HNO₃ at 14 kg Nha^–1 yr^–1. Litter decompositionrates of Fagus grandifolia and Acerrubrum were substantially higher at WL andPHC compared to HF but were not affected byexperimental N additions. Results usingmineral soil bags showed no effects of Naddition on N and C concentrations in soilorganic matter, but C and N concentrationincreases were less at WL and PHC compared toHF. Soil solution nitrate (NO₃ ^–)concentrations at 15-cm depth in the referenceplots were higher at PHC than at WL and HFwhile at 50-cm concentrations were higher atPHC and WL than at HF. The reference plots atthe two sites (WL and PHC) with the highestatmospheric inputs of N exhibited lower Nretention (53 and 33%, respectively) than HF(68%) in reference plots. The greatestincrease in NO₃ ^– loss in response tothe experimental treatments occurred at HFwhere the HNO₃ additions resulted in thehighest NO₃ ^– concentrations andlowest N retentions. In contrast, at WL andPHC increases in soil water NO₃ ^–were not evident in response to experimental Nadditions. The results suggest that the twosites (WL and PHC) in the western Adirondacksdid not respond to additional N inputsalthough they have experienced elevatedatmospheric N inputs and higher N drainagelosses in reference plots than the HF site inthe central Adirondacks. Some of thesedifferences in site response may have alsobeen a function of stand age of WL and PHCthat were younger (24 and 33 years,respectively) than the HF (age 70).Highest NO₃ ^– fluxes in thereference plots across the sites correspondedto higher ¹⁵N values in soil andplants. An experimental addition experimentat PHC found that the forest floor and themineral soil were the largest sinks forexperimentally added N.     

Net nitrogen mineralization in soils under six indigenous tree species,an abandoned pasture and a secondary forest in the atlantic lowlands of costa rica

Nitrogen mineralization, nitrification potentials, pH, total N, C, extractable P and cations were measured in soils under 4-year-old, mono-specific stands of six fast-growing, native tree species, an abandoned pasture, and a 20-year-old secondary forest, as part of a study on the use of indigenous tree species for rehabilitation of soil fertility on degraded pastures at the La Selva Biological Station in the Atlantic humid lowlands of Costa Rica. Soil net nitrification potential rates were higher under two N-fixing, leguminous species,Stryphnodendron microstachyum Poepp. et Endl. (1.1–1.9 mg kg^–1 day^–1) andDalbergia tucurensis Donn. Smith (0.7–1.5 mg kg^–1 day^–1), than under the non-N-fixing trees in the plantation,Vochysia guatemalesis Don. Sm.,Vochysia ferruginea Mart,Dipteryx panamensis (Pittier) Record and Mell andHyeronima alchorneoides Fr. Allemao (0.2–0.8 mg kg^–1 day^–1). Values under the N-fixing trees were comparable to those found in secondary forest. There were no statistically significant differences in soil total N or in other nurtients between the species. Results of pH measurements done before and after incubation did not show any clear evidence of a pH drop attributable to nitrification.     

Forests losing large quantities of nitrogen have elevated 15N:14N ratios

Summary Urea (U) and ammonium nitrate (AN) had been applied to a Scots pine (Pinus sylvestris L.) forest in northern Sweden for 18 consecutive years at four doses resulting in total N applications ranging from 0 to 1980 kg ha^–1. The ¹⁵N abundance ( ¹⁵N) of the grass Deschampsia flexuosa (L.) Trin. increased linearly (from –0.7 to 11.0) with application rate in the case of U. The response to AN was in the same direction but smaller. While others have shown that the initial response of nitrogen-limited systems to additions of N is a change of ¹⁵N abundance towards that of added N, this study shows that further and excessive additions leads to a retention of ¹⁵N. Monitoring ¹⁵N abundance over time in dose-response trials of this type thus opens new possibilities to estimate critical loads of N and the point of nitrogen saturation.     

Stable soil nitrogen accumulation and flexible organic matter stoichiometry during primary floodplain succession

Large increases in nitrogen (N) inputs to terrestrial ecosystems typically have small effects on immediate N outputs because most N is sequestered in soil organic matter. We hypothesized that soil organic N storage and the asynchrony between N inputs and outputs result from rapid accumulation of N in stable soil organic pools. We used a successional sequence on floodplains of the Tanana River near Fairbanks, Alaska to assess rates of stable N accumulation in soils ranging from 1 to 500+ years old. One-year laboratory incubations with repeated leaching separated total soil N into labile (defined as inorganic N leached) and stable (defined as total minus labile N) pools. Stable N pools increased faster (2 g N m^–2 yr^–1) than labile N (0.4 g N m^–2 yr^–1) pools during the first 50 years of primary succession; labile N then plateaued while stable and total N continued to increase. Soil C pools showed similar trends, and stable N was correlated with stable C (r² = 0.95). From 84 to 95 % of soil N was stable during our incubations. Over successional time, the labile N pool declined as a proportion of total N, but remained large on an aerial basis (up to 38 g N m^–2). The stoichiometry of stable soil N changed over successional time; C:N ratios increased from 10 to 22 over 275 years (r² = 0.69). A laboratory ¹⁵N addition experiment showed that soils had the capacity to retain much more N than accumulated naturally during succession. Our results suggest that most soil N is retained in a stable organic pool that can accumulate rapidly but is not readily accessible to microbial mineralization. Because stable soil organic matter and total ecosystem organic matter have flexible stoichiometry, net ecosystem production may be a poor predictor of N retention on annual time scales.     

Nitrogen fixation by white clover in pastures grazed by dairy cows: Temporal variation and effects of nitrogen fertilization

Effects of rate of nitrogen (N) fertilizer and stocking rate on production and N₂ fixation by white clover (Trifolium repens L.) grown with perennial ryegrass (Lolium perenne L.) were determined over 5 years in farmlets near Hamilton, New Zealand. Three farmlets carried 3.3 dairy cows ha^–1 and received urea at 0, 200 or 400 kg N ha^–1 yr^–1 in 8–10 split applications. A fourth farmlet received 400 kg N ha^–1 yr^–1 and had 4.4 cows ha^–1.There was large variation in annual clover production and total N₂ fixation, which in the 0 N treatment ranged from 9 to 20% clover content in pasture and from 79 to 212 kg N fixed ha^–1 yr^–1. Despite this variation, total pasture production in the 0 N treatment remained at 75–85% of that in the 400 N treatments in all years, due in part to the moderating effect of carry-over of fixed N between years.Fertilizer N application decreased the average proportion of clover N derived from N₂ fixation (P_N; estimated by ¹⁵N dilution) from 77% in the 0 N treatment to 43–48% in the 400 N treatments. The corresponding average total N₂ fixation decreased from 154 kg N ha^–1 yr^–1 to 39–53 kg N ha^–1 yr^–1. This includes N₂ fixation in clover tissue below grazing height estimated at 70% of N₂ fixation in above grazing height tissue, based on associated measurements, and confirmed by field N balance calculations. Effects of N fertilizer on clover growth and N₂ fixation were greatest in spring and summer. In autumn, the 200 N treatment grew more clover than the 0 N treatment and N₂ fixation was the same. This was attributed to more severe grazing during summer in the 0 N treatment, resulting in higher surface soil temperatures and a deleterious effect on clover stolons.In the 400 N treatments, a 33% increase in cow stocking rate tended to decrease P_N from 48 to 43% due to more N cycling in excreta, but resulted in up to 2-fold more clover dry matter and N₂ fixation because lower pasture mass reduced grass competition, particularly during spring.     

Natural 15N abundances of maize and soil amended with urea and composted pig manure

To investigate the effect of inorganic fertilizer and composted manure amendments on the N isotope composition (delta ¹⁵N) of crop and soil, maize (Zea mays L.) was cultivated under greenhouse conditions for 30, 40, 50, 60, and 70 days. Composted pig manure (delta ¹⁵N= +13.9) and urea (-2.3) were applied at 0 and 0 kg N ha^–1 (C0U0), 0 and 150 kg N ha^–1 (C0U2), 150 and 0 kg N ha^–1 (C2U0), and 75 and 75 kg N ha^–1 (C1U1), respectively. The delta ¹⁵N of total soil-N was not affected by both amendments, but delta ¹⁵N of NH⁺ ₄ and NO^– ₃ provided some information on the N isotope fractionation in soil. During the early growth stage, significant differences (P < 0.05) in delta ¹⁵N among maize subjected to different treatments were observed. After 30 days of growth, the delta ¹⁵N values of maize were +6.6 for C0U0, +1.1 for C0U2, +7.7 for C2U0, and +4.5 for C1U1. However, effects of urea and composted manure application on maize delta ¹⁵N progressively decreased with increasing growth period, probably due to isotope fractionation accompanying N losses and increased uptake of soil-derived N by maize. After 70 days of growth, delta ¹⁵N of leaves and grains of maize amended with composted pig manure were significantly (P < 0.05) higher than those with urea. The temporal variations in delta ¹⁵N of maize amended with urea and composted manure indicate that plant delta ¹⁵N is generally not a good tracer for N sources applied to field. Our data can be used in validation of delta ¹⁵N fractionation models in relation to N source inputs.