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.     

Immobilization, stabilization and remobilization of nitrogen in forest soils at elevated CO2: a 15N and 13C tracer study

The fate of immobilized N in soils is one of the great uncertainties in predicting C sequestration at increased CO₂ and N deposition. In a dual isotope tracer experiment (¹³C, ¹⁵N) within a 4‐year CO₂ enrichment (+200 ppm_v) study with forest model ecosystems, we (i) quantified the effects of elevated CO₂ on the partitioning of N; (ii) traced immobilized N into physically separated pools of soil organic matter (SOM) with turnover rates known from their ¹³C signals; and (iii) estimated the remobilization and thus, the bio‐availability of newly sequestered C and N. (1) CO₂ enrichment significantly decreased NO₃^? concentrations in soil waters and export from 1.5 m deep lysimeters by 30–80%. Consequently, elevated CO₂ increased the overall retention of N in the model ecosystems. (2) About 60–80% of added ¹⁵NH₄¹⁵NO₃ were retained in soils. The clay fraction was the greatest sink for the immobilized ¹⁵N sequestering 50–60% of the total new soil N. SOM associated with clay contained only 25% of the total new soil C pool and had small C/N ratios (<13), indicating that it consists of humified organic matter with a relatively slow turn over rate. This implies that added ¹⁵N was mainly immobilized in stable mineral‐bound SOM pools. (3) Incubation of soils for 1 year showed that the remobilization of newly sequestered N was three to nine times smaller than that of newly sequestered C. Thus, inorganic inputs of N were stabilized more effectively in soils than C. Significantly less newly sequestered N was remobilized from soils previously exposed to elevated CO₂. In summary, our results show firstly that a large fraction of inorganic N inputs becomes effectively immobilized in relative stable SOM pools and secondly that elevated CO₂ can increase N retention in soils and hence it may tighten N cycling and diminish the risk of nitrate leaching to groundwater.     

Influence of Tree Species on Forest Nitrogen Retention in the Catskill Mountains, New York, USA

This study examines the effect of four tree species on nitrogen (N) retention within forested catchments of the Catskill Mountains, New York (NY). We conducted a 300-day ¹⁵N field tracer experiment to determine how N moves through soil, microbial, and plant pools under different tree species and fertilization regimes. Samples were collected from single-species plots of American beech (Fagus grandifolia Ehrh.), eastern hemlock (Tsuga canadensis L.), red oak (Quercus rubra L.), and sugar maple (Acer saccharum Marsh). Using paired plots we compared the effects of ambient levels of N inputs (11 kg N/ha/y) to additions of 50 kg N/ha/y that began 1.5 years prior to and continued throughout this experiment. Total plot ¹⁵N recovery (litter layer, organic and mineral soil to 12 cm, fine roots, and aboveground biomass) did not vary significantly among tree species, but the distribution of sinks for ¹⁵N within the forest ecosystem did vary. Recovery in the forest floor was significantly lower in sugar maple stands compared to the other species. ¹⁵Nitrogen recovery was 22% lower in the fertilized plots compared to the ambient plots and red oak stands had the largest drop in ¹⁵N recovery as a result of N fertilization. Aboveground biomass became a significantly greater ¹⁵N sink with fertilization, although it retained less than 1% of the tracer addition. These results indicate that different forest types vary in the amount of N retention in the forest floor, and that forest N retention may change depending upon N inputs.     

Detrital Controls on Soil Solution N and Dissolved Organic Matter in Soils: A Field Experiment

We established a long-term field study in an old growth coniferous forest at the H.J. Andrews Experimental Forest, OR, USA, to address how detrital quality and quantity control soil organic matter accumulation and stabilization. The Detritus Input and Removal Treatments (DIRT) plots consist of treatments that double leaf litter, double woody debris inputs, exclude litter inputs, or remove root inputs via trenching. We measured changes in soil solution chemistry with depth, and conducted long-term incubations of bulk soils from different treatments in order to elucidate effects of detrital inputs on the relative amounts and lability of different soil C pools. In the field, the addition of woody debris increased dissolved organic carbon (DOC) concentrations in O-horizon leachate and at 30 cm, but not at 100 cm, compared to control plots, suggesting increased rates of DOC retention with added woody debris. DOC concentrations decreased through the soil profile in all plots to a greater degree than did dissolved organic nitrogen (DON), most likely due to preferential sorption of high C:N hydrophobic dissolved organic matter (DOM) in upper horizons; percent hydrophobic DOM decreased significantly with depth, and hydrophilic DOM had a much lower and less variable C:N ratio. Although laboratory extracts of different litter types showed differences in DOM chemistry, percent hydrophobic DOM did not differ among soil solutions from different detrital treatments in the field, suggesting that microbial processing of DOM leachate in the field consumed easily degradable components, thus equalizing leachate chemistry among treatments. Total dissolved N leaching from plots with intact roots was very low (0.17 g m⁻² year⁻¹), slightly less than measured deposition to this very unpolluted forest (~s 0.2 g m⁻² year⁻¹). Total dissolved N losses showed significant increases in the two treatments without roots whereas concentrations of DOC decreased. In these plots, N losses were less than half of estimated plant uptake, suggesting that other mechanisms, such as increased microbial immobilization of N, accounted for retention of N in deep soils. In long-term laboratory incubations, soils from plots that had both above- and below-ground litter inputs excluded for 5 years showed a trend towards lower DOC loss rates, but not lower respiration rates. Soils from plots with added wood had similar respiration and DOC loss rates as control soils, suggesting that the additional DOC sorption observed in the field in these soils was stabilized in the soil and not readily lost upon incubation.     

Incorporation of Plant Residues into Soil Organic Matter Fractions With Grassland Management Practices in the North American Midwest

Disturbed grassland soils are often cited as having the potential to store large amounts of carbon (C). Fertilization of grasslands can promote soil C storage, but little is known about the generation of recalcitrant pools of soil organic matter (SOM) with management treatments, which is critical for long-term soil C storage. We used a combination of soil incubations, size fractionation and acid hydrolysis of SOM, [C], [N], and stable isotopic analyses, and biomass quality indices to examine how fertilization and haying can impact SOM dynamics in Kansan grassland soils. Fertilized soils possessed 113% of the C possessed by soils subjected to other treatments, an increase predominantly harbored in the largest size fraction (212–2,000 μm). This fraction is frequently associated with more labile material. Haying and fertilization/haying, treatments that more accurately mimic true management techniques, did not induce any increase in soil C. The difference in ¹⁵N-enrichment between size fractions was consistent with a decoupling of SOM processing between pools with fertilization, congruent with gains of SOM in the largest size fraction promoted by fertilization not moving readily into smaller fractions that frequently harbor more recalcitrant material. Litterfall and root biomass C inputs increased 104% with fertilization over control plots, and this material possessed lower C:N ratios. Models of incubation mineralization kinetics indicate that fertilized soils have larger pools of labile organic C. Model estimates of turnover rates of the labile and recalcitrant C pools did not differ between treatments (65.5 ± 7.2 and 2.9 ± 0.3 μg C d⁻¹, respectively). Although fertilization may promote greater organic inputs into these soils, much of that material is transformed into relatively labile forms of soil C; these data highlight the challenges of managing grasslands for long-term soil C sequestration.     

Linking Carbon Saturation Concepts to Nitrogen Saturation and Retention

Recent advances in soil C saturation concepts have increased our understanding of soil C storage and mineralization without explicit links to N retention and saturation theories. Here, we exploit soil texture and organic matter (OM) gradients in a Maryland, USA hardwood forest to test hypotheses that link soil organic C saturation with soil ¹⁵N retention and nitrification. At our site, mineral-associated OM (MAOM) N concentrations in the silt + clay particle fraction (g MAOM-N g silt + clay⁻¹) were negatively correlated with the fraction of NH₄-N transferred to MAOM during a 3-day in situ incubation (R = −0.85), but positively correlated with potential net nitrification (R = 0.76). Moreover, the fraction of NH₄-N transferred to MAOM was negatively correlated with potential net nitrification (R = −0.76). Due to physico-chemical stabilization mechanisms, MAOM is considered to be resistant to mineralization. Carbon saturation theory suggests that the proportion of new C inputs that can be stabilized in MAOM decreases in proportion to the amount of C already present in the fraction; C inputs not stabilized in MAOM are susceptible to rapid mineralization. We demonstrate that NH₄-N stabilization in MAOM is similar to C stabilization in MAOM and associated with nitrification, thereby extending soil C saturation theory to mineral N and linking it with N retention and saturation theories. These data and concepts complement N saturation models that emphasize vegetation type, N input levels, and microbial turnover. Incorporating the OM retention capacity of fine mineral particles into N saturation theory can improve predictions of N saturation rates and resolve inconsistent relationships between soil organic matter, texture, N mineralization, and N retention.     

Decomposition nitrogen is better retained than simulated deposition from mineral amendments in a temperate forest

Nitrogen (N) deposition (N_DEP) drives forest carbon (C) sequestration but the size of this effect is still uncertain. In the field, an estimate of these effects can be obtained by applying mineral N fertilizers over the soil or forest canopy. A ¹⁵N label in the fertilizer can be then used to trace the movement of the added N into ecosystem pools and deduce a C effect. However, N recycling via litter decomposition provides most of the nutrition for trees, even under heavy N_DEP inputs. If this recycled litter nitrogen is retained in ecosystem pools differently to added mineral N, then estimates of the effects of N_DEP on the relative change in C (?C/?N) based on short‐term isotope‐labelled mineral fertilizer additions should be questioned. We used ¹⁵N labelled litter to track decomposed N in the soil system (litter, soils, microbes, and roots) over 18 months in a Sitka spruce plantation and directly compared the fate of this ¹⁵N to an equivalent amount in simulated N_DEP treatments. By the end of the experiment, three times as much ¹⁵N was retained in the O and A soil layers when N was derived from litter decomposition than from mineral N additions (60% and 20%, respectively), primarily because of increased recovery in the O layer. Roots expressed slightly more ¹⁵N tracer from litter decomposition than from simulated mineral N_DEP (7.5% and 4.5%) and compared to soil recovery, expressed proportionally more ¹⁵N in the A layer than the O layer, potentially indicating uptake of organic N from decomposition. These results suggest effects of N_DEP on forest ?C/?N may not be apparent from mineral ¹⁵N tracer experiments alone. Given the importance of N recycling, an important but underestimated effect of N_DEP is its influence on the rate of N release from litter.     

Changes in Soil Carbon and Nitrogen after Contrasting Land-use Transitions in Northeastern Costa Rica

Northeastern Costa Rica is a mosaic of primary and secondary forests, tree plantations, pastures, and cash crops. Many studies have quantified the effects of one type of land-use transition (for example, deforestation or reforestation) on soil properties such as organic carbon (C) storage, but few have compared different land-use transitions simultaneously. We can best understand the effects of land-use change on regional and global ecosystem processes by considering all of the land-use transitions that occur in a landscape. In this study, I examined the changes in total soil C and nitrogen (N) pools (to 0.3 m) that have accompanied different land-use transitions in a 140,000-ha region in northeastern Costa Rica. I paired sites that had similar topography and soils but differed in recent land-use history. The following land-use transitions were represented: 12 conversions of primary forests to banana plantations, 15 conversions of pastures to cash crops, and four conversions of pastures to Vochysia guatemalensis tree plantations. The conversion of forests to bananas decreased soil C concentrations and inventories (Mg C ha^–1) in the surface soil by 37% and 16.5%, respectively. The conversion of pastures to cash crops reduced soil C concentrations and inventories to the same extent that forest-to-banana cropping did. Furthermore, young Vochysia plantations do not appear to increase soil C storage, at least over the 1st decade. When data from all land-use transitions were pooled, the difference in root biomass and leaf litter pools between land-use pairs explained 50% of the differences in soil C concentrations and 36% of the differences in soil C inventories. Thus, reduced productivity or C inputs to the soil is one mechanism that could explain the losses in soil C pools with land-use change. In this landscape, losses of soil C due to cultivation are rapid, whereas re- accumulation rates are slow. Total soil N pools (0–10 cm) were also reduced after the conversion of forests to banana plantations or the conversion of pastures to crops, despite fertilization of the cropped soils. This suggests that the added N fertilizer is not retained but instead is exported via produce, N gas emissions, and hydrologic processes.     

5300-Year-old soil carbon is less primed than young soil organic matter

Soils harbor more than three times as much carbon (C) as the atmosphere, a large fraction of which (stable organic matter) serves as the most important global C reservoir due to its long residence time. Litter and root inputs bring fresh organic matter (FOM) into the soil and accelerate the turnover of stable C pools, and this phenomenon is termed the “priming effect” (PE). Compared with knowledge about labile soil C pools, very little is known about the vulnerability of stable C to priming. Using two soils that substantially differed in age (500 and 5300 years before present) and in the degree of chemical recalcitrance and physical protection of soil organic matter (SOM), we showed that leaf litter amendment primed 264% more organic C from the young SOM than from the old soil with very stable C. Hierarchical partitioning analysis confirmed that SOM stability, reflected mainly by available C and aggregate protection of SOM, is the most important predictor of leaf litter-induced PE. The addition of complex FOM (i.e., leaf litter) caused a higher bacterial oligotroph/copiotroph (K-/r-strategists) ratio, leading to a PE that was 583% and 126% greater than when simple FOM (i.e., glucose) was added to the young and old soils, respectively. This implies that the PE intensity depends on the chemical similarity between the primer (here FOM) and SOM. Nitrogen (N) mining existed when N and simple FOM were added (i.e., Glucose+N), and N addition raised the leaf litter-induced PE in the old soil that had low N availability, which was well explained by the microbial stoichiometry. In conclusion, the PE induced by FOM inputs strongly decreases with increasing SOM stability. However, the contribution of stable SOM to CO₂ efflux cannot be disregarded due to its huge pool size.     

The impact of long-term elevated CO2 on C and N retention in stable SOM pools

Elevated atmospheric CO₂ frequently increases plant production and concomitant soil C inputs, which may cause additional soil C sequestration. However, whether the increase in plant production and additional soil C sequestration under elevated CO₂ can be sustained in the long-term is unclear. One approach to study C–N interactions under elevated CO₂ is provided by a theoretical framework that centers on the concept of progressive nitrogen limitation (PNL). The PNL concept hinges on the idea that N becomes less available with time under elevated CO₂. One possible mechanism underlying this reduction in N availability is that N is retained in long-lived soil organic matter (SOM), thereby limiting plant production and the potential for soil C sequestration. The long-term nature of the PNL concept necessitates the testing of mechanisms in field experiments exposed to elevated CO₂ over long periods of time. The impact of elevated CO₂ and ¹⁵N fertilization on L. perenne and T. repens monocultures has been studied in the Swiss FACE experiment for ten consecutive years. We applied a biological fractionation technique using long-term incubations with repetitive leaching to determine how elevated CO₂ affects the accumulation of N and C into more stable SOM pools. Elevated CO₂ significantly stimulated retention of fertilizer-N in the stable pools of the soils covered with L. perenne receiving low and high N fertilization rates by 18 and 22%, respectively, and by 45% in the soils covered by T. repens receiving the low N fertilization rate. However, elevated CO₂ did not significantly increase stable soil C formation. The increase in N retention under elevated CO₂ provides direct evidence that elevated CO₂ increases stable N formation as proposed by the PNL concept. In the Swiss FACE experiment, however, plant production increased under elevated CO₂, indicating that the additional N supply through fertilization prohibited PNL for plant production at this site. Therefore, it remains unresolved why elevated CO₂ did not increase labile and stable C accumulation in these systems.     

Greater Soil Carbon Sequestration under Nitrogen-fixing Trees Compared with <Emphasis Type="Italic">Eucalyptus</Emphasis> Species

Forests with nitrogen-fixing trees (N–fixers) typically accumulate more carbon (C) in soils than similar forests without N–fixing trees. This difference may develop from fundamentally different processes, with either greater accumulation of recently fixed C or reduced decomposition of older soil C. We compared the soil C pools under N–fixers with Eucalyptus (non–N–fixers) at four tropical sites: two sites on Andisol soils in Hawaii and two sites on Vertisol and Entisol soils in Puerto Rico. Using stable carbon isotope techniques, we tracked the loss of the old soil organic C from the previous C₄ land use (SOC₄) and the gain of new soil organic C from the C₃, N–fixer, and non–N–fixer plantations (SOC₃). Soils beneath N–fixing trees sequestered 0.11 ± 0.07 kg m⁻² y⁻¹ (mean ± one standard error) of total soil organic carbon (SOC_T) compared with no change under Eucalyptus (0.00 ± 0.07 kg m⁻² y⁻¹; P = 0.02). About 55% of the greater SOC_T sequestration under the N–fixers resulted from greater retention of old SOC₄, and 45% resulted from greater accretion of new SOC₃. Soil N accretion under the N–fixers explained 62% of the variability of the greater retention of old SOC₄ under the N–fixers. The greater retention of older soil C under N–fixing trees is a novel finding and may be important for strategies that use reforestation or afforestation to offset C emissions. Received 12 March 2001; accepted 5 October 2001.     

Litter type control on soil C and N stabilization dynamics in a temperate forest

While plant litters are the main source of soil organic matter (SOM) in forests, the controllers and pathways to stable SOM formation remain unclear. Here, we address how litter type (¹³C/¹⁵N‐labeled needles vs. fine roots) and placement‐depth (O vs. A horizon) affect in situ C and N dynamics in a temperate forest soil after 5 years. Litter type rather than placement‐depth controlled soil C and N retention after 5 years in situ, with belowground fine root inputs greatly enhancing soil C (x1.4) and N (x1.2) retention compared with aboveground needles. While the proportions of added needle and fine root‐derived C and N recovered into stable SOM fractions were similar, they followed different transformation pathways into stable SOM fractions: fine root transfer was slower than for needles, but proportionally more of the remaining needle‐derived C and N was transferred into stable SOM fractions. The stoichiometry of litter‐derived C vs. N within individual SOM fractions revealed the presence at least two pools of different turnover times (per SOM fraction) and emphasized the role of N‐rich compounds for long‐term persistence. Finally, a regression approach suggested that models may underestimate soil C retention from litter with fast decomposition rates.     

Plant and soil natural abundance <Emphasis Type="Italic">δ</Emphasis><Superscript>15</Superscript>N: indicators of relative rates of nitrogen cycling in temperate forest ecosystems

Watersheds within the Catskill Mountains, New York, receive among the highest rates of nitrogen (N) deposition in the northeastern United States and are beginning to show signs of N saturation. Despite similar amounts of N deposition across watersheds within the Catskill Mountains, rates of soil N cycling and N retention vary significantly among stands of different tree species. We examined the potential use of δ ¹⁵N of plants and soils as an indicator of relative forest soil N cycling rates. We analyzed the δ ¹⁵N of foliage, litterfall, bole wood, surface litter layer, fine roots and organic soil from single-species stands of American beech (Fagus grandifolia), eastern hemlock (Tsuga canadensis), red oak (Quercus rubra), and sugar maple (Acer saccharum). Fine root and organic soil δ ¹⁵N values were highest within sugar maple stands, which correlated significantly with higher rates of net mineralization and nitrification. Results from this study suggest that fine root and organic soil δ ¹⁵N can be used as an indicator of relative rates of soil N cycling. Although not statistically significant, δ ¹⁵N was highest within foliage, wood and litterfall of beech stands, a tree species associated with intermediate levels of soil N cycling rates and forest N retention. Our results show that belowground δ ¹⁵N values are a better indicator of relative rates of soil N cycling than are aboveground δ ¹⁵N values.     

Accumulation of Carbon and Nitrogen in Residential Soils with Different Land-Use Histories

Urban areas are growing in size and importance; however, we are only beginning to understand how the process of urbanization influences ecosystem dynamics. In particular, there have been few assessments of how the land-use history and age of residential soils influence carbon (C) and nitrogen (N) pools and fluxes, especially at depth. In this study, we used 1-m soil cores to evaluate soil profile characteristics and C and N pools in 32 residential home lawns that differed by previous land use and age, but had similar soil types. These were compared to soils from eight forested reference sites. Residential soils had significantly higher C and N densities than nearby forested soils of similar types (6.95 vs. 5.44 kg C/m² and 552 vs. 403 g N/m², P < 0.05). Results from our chronosequence suggest that soils at residential sites that were previously in agriculture have the potential to accumulate C (0.082 kg C/m²/y) and N (8.3 g N/m²/y) rapidly after residential development. Rates of N accumulation at these sites were similar in magnitude to estimated fertilizer N inputs, confirming a high capacity for N retention. Residential sites that were forested prior to development had higher C and N densities than present-day forests, but our chronosequence did not reveal a significant pattern of increasing C and N density over time in previously forested sites. These data suggest that soils in residential areas on former agricultural land have a significant capacity to sequester C and N. Given the large area of these soils, they are undoubtedly significant in regional C and N balances.     

Long-Term Nitrogen Additions and Nitrogen Saturation in Two Temperate Forests

This article reports responses of two different forest ecosystems to 9 years (1988–96) of chronic nitrogen (N) additions at the Harvard Forest, Petersham, Massachusetts. Ammonium nitrate (NH₄NO₃) was applied to a pine plantation and a native deciduous broad-leaved (hardwood) forest in six equal monthly doses (May–September) at four rates: control (no fertilizer addition), low N (5 g N m^-2 y^-1), high N (15 g N m^-2 y^-1), and low N + sulfur (5 g N m^-2 y^-1 plus 7.4 g S m^-2 y^-1). Measurements were made of net N mineralization, net nitrification, N retention, wood production, foliar N content and litter production, soil C and N content, and concentrations of dissolved organic carbon (DOC) and nitrogen (DON) in soil water. In the pine stand, nitrate losses were measured after the first year of additions (1989) in the high N plot and increased again in 1995 and 1996. The hardwood stand showed no significant increases in nitrate leaching until 1995 (high N only), with further increases in 1996. Overall N retention efficiency (percentage of added N retained) over the 9-year period was 97–100% in the control and low N plots of both stands, 96% in the hardwood high N plot, and 85% in the pine high N plot. Storage in aboveground biomass, fine roots, and soil extractable pools accounted for only 16–32% of the added N retained in the amended plots, suggesting that the one major unmeasured pool, soil organic matter, contains the remaining 68–84%. Short-term redistribution of ¹⁵N tracer at natural abundance levels showed similar division between plant and soil pools. Direct measurements of changes in total soil C and N pools were inconclusive due to high variation in both stands. Woody biomass production increased in the hardwood high N plot but was significantly reduced in the pine high N plot, relative to controls. A drought-induced increase in foliar litterfall in the pine stand in 1995 is one possible factor leading to a measured increase in N mineralization, nitrification, and nitrate loss in the pine high N plot in 1996. Received 2 April 1999; Accepted 29 October 1999.     

Soil amino-acid availability across a temperate-forest fertility gradient

Despite increasing recognition that free amino acids can be an important source of N for plant uptake, we have a poor understanding of environmental variation in the availability of amino-acid N in soils outside of arctic, alpine and boreal regions. I investigated patterns of amino-acid availability along a temperate forest fertility gradient ranging from low mineral N availability, oak-dominated forests to high mineral N availability, maple-basswood forests (5 sites). I measured standing pools of free amino acids, soluble peptides, ammonium and nitrate, rates of amino acid production (native proteolysis activity) and rates of consumption of a ¹⁵N-labeled leucine tracer. Standing pools of amino acid N decreased consistently along the fertility gradient from the low fertility black oak/white oak system to the high fertility sugar maple/basswood system, with a 25-fold difference in pool sizes between the poorest and richest sites. Standing pools of soluble peptides varied little among sites, instead, the relationship between free amino acids and peptides changed markedly across the gradient. At low fertility sites free amino acids were positively correlated with soluble peptides, whereas free amino acid pools were universally low at high fertility sites, regardless of peptide pools. Assays for native proteolysis activity indicated that amino acid production did not vary significantly among sites. Recovery of leucine tracer in inorganic (NH₄ ⁺ and NO₃ ⁻) pools and in residual soil organic matter both increased with increasing soil fertility; however, total consumption of the added amino-acid tracer did not vary among sites. Results from this study demonstrate that free amino acids can make an important contribution to potentially plant-available N pools in temperate forest soils, particularly at low fertility sites.     

Long-term movement of 15N tracers into fine woody debris under chronically elevated N inputs

Two key questions in the study of large-scale C (carbon) and N (nitrogen) cycling in temperate forests are how N cycling in soil detritus controls ecosystem-level retention of elevated N deposition, and whether elevated N deposition is likely to cause increases in C pools. The large C:N ratios in woody detritus make it a potentially important contributor to N retention, if N immobilization increases, and a potentially important contributor to C sequestration, if pool sizes increase. We studied N concentrations, C:N ratios, and pool sizes of N and biomass in fine woody debris (FWD < 5 cm diam.) 12 years into a long-term N-amendment study in two contrasting forests, a naturally-regenerated forest dominated by Quercus spp., and a 63-yr old plantation of Pinus resinosa. We also quantitatively recovered ¹⁵N tracers (originally applied as ¹⁵NH₄ and ¹⁵NO₃) in FWD, eight years following their application in the same study, in both ambient and N-amended plots. We used these data to test predictions of tracer redistributions made by a biogeochemical process model that included ¹⁵N. Results from the N pool-size analysis and the ¹⁵N tracer-recovery analysis indicated that under elevated N inputs of 5 g N m^–2 yr^–1 (as NH₄NO₃) over the decadal time period, only 0.15%–0.76% of the elevated N inputs were recovered in FWD of N-amended plots relative to ambient. Any increase in N immobilization in wood appeared to be minimal, in agreement with model predictions. Under N amendments, pool sizes of C in FWD were not significantly different from ambient, whereas pool sizes of N were marginally higher. Patterns of ¹⁵NH₄ vs. ¹⁵NO₃ recovery, treatment differences, and forest-type differences suggested that plant uptake, rather than detrital immobilization, was the dominant mechanism of ¹⁵N tracer movement into FWD. This result indicates that plant-soil cycling operating over a decadal time scale or longer controls C:N ratios and N pool sizes in woody debris.     

Carbon and nitrogen in a temperate agroforestry system: Using stable isotopes as a tool to understand soil dynamics

Natural exchanges of carbon (C) between the atmosphere, the oceans, and terrestrial ecosystems are currently being modified through human activities as a result of fossil fuel burning and the conversion of tropical forests to agricultural land. These activities have led to a steady increase of atmospheric carbon dioxide (CO₂) over the last two Centuries. The goal of this study was to determine the potential of temperate agroforestry systems to sequester C in soil. Therefore, changes in the soil organic C (SOC) and nitrogen (N) pools were quantified and the δ¹³C and δ¹⁵N stable isotope technique was applied to assess soil C and N dynamics in a 13-year old hybrid poplar alley cropping system in Southern Canada. Results from this study showed that after 13 years of alley cropping the SOC and N pools did not differ significantly (p = 0.01) with distance from the tree row, although a trend of a larger SOC and N pool near the tree row could be observed. Soil organic C after 13 years of alley cropping, was 19 mg C g⁻¹ compared to 11 mg C g⁻¹ upon initiation of agroforestry. Soil organic C and N were not evenly distributed throughout the plow layer. The largest C and N pool occurred in the top 20 cm, which is due to the accumulation of organic material in the upper horizons as a result of no-till cultivation. The entire soil, to a 40 cm depth, showed a δ¹³C shift to that of C₃ residue. This shift reflects the greater input of residues from C₃ plants such as that derived from beans, wheat, and hybrid poplar leaf litterfall. The proportion of C derived from a C₃ source ranged from 64 to 69% to a 40 cm depth. The soil δ¹⁵N signature of this study is similar to that of mineral soil, and reflect values characteristic of N mineralization processes. However, the entire soil shows a positive shift in δ¹⁵N as a result of historical additions of manure and current use of mineral fertilizers, and ongoing processes of denitrification and nitrate leaching, which leads to an enrichment of the soil.     

Soluble Organic Nitrogen Pools in Forest soils of Subtropical Australia

Soil soluble organic N (SON) plays an important role in N biogeochemical cycling. In this study, 22 surface forest soils (0–10 cm) were collected from southeast Queensland, Australia, to investigate the size of SON pools extracted by water and salt solutions. Approximately 5–45 mg SON kg⁻¹, 2–42 mg SON kg⁻¹ and 1–24 SON mg kg⁻¹ were extracted by 2 M KCl, 0.5 M K₂SO₄ and water, on average, corresponding to about 21.1, 13.5 and 7.0 kg SON ha⁻¹ at the 0–10 cm forest soils, respectively. These SON pools, on average, accounted for 39% (KCl extracts), 42% (K₂SO₄ extracts) and 43% (water extracts) of total soluble N (TSN), and 2.3% (KCl extracts), 1.3% (K₂SO₄ extracts) and 0.7% (water extracts) of soil total N, respectively. Large variation in SON pools observed across the sites in the present study may be attributed to a combination of factors including soil types, tree species, management practices and environmental conditions. Significant relationships were observed among the SON pools extracted by water, KCl and K₂SO₄ and microbial biomass N (MBN). In general, KCl and K₂SO₄ extracted more SON than water from the forest soils, while KCl extracted more SON than K₂SO₄. The SON and soluble organic C (SOC) in KCl, K₂SO₄ and water extracts were all positively related to soil organic C, total N and clay contents. This indicates that clay and soil organic matter play a key role in the retention of SON in soil.     

Multiyear fate of a 15N tracer in a mixed deciduous forest: retention,redistribution, and differences by mycorrhizal association

The impact of atmospheric nitrogen deposition on forest ecosystems depends in large part on its fate. Past tracer studies show that litter and soils dominate the short‐term fate of added ¹⁵N, yet few have examined its longer term dynamics or differences among forest types. This study examined the fate of a ¹⁵N‐ tracer over 5–6 years in a mixed deciduous stand that was evenly composed of trees with ectomycorrhizal and arbuscular mycorrhizal associations. The tracer was expected to slowly mineralize from its main initial fate in litter and surface soil, with some ¹⁵N moving to trees, some to deeper soil, and some net losses. Recovery of added ¹⁵N in trees and litterfall totaled 11.3% both 1 and 5–6 years after the tracer addition, as ¹⁵N redistributed from fine and especially coarse roots into cumulative litterfall and small accumulations in woody tissues. Estimates of potential carbon sequestration from tree ¹⁵N recovery amounted to 12–14 kg C per kg of N deposition. Tree ¹⁵N acquisition occurred within the first year after the tracer addition, with no subsequent additional net transfer of ¹⁵N from detrital to plant pools. In both years, ectomycorrhizal trees gained 50% more of the tracer than did trees with arbuscular mycorrhizae. Much of the ¹⁵N recovered in wood occurred in tree rings formed prior to the ¹⁵N addition, demonstrating the mobility of N in wood. Tracer recovery rapidly decreased over time in surface litter material and accumulated in both shallow and deep soil, perhaps through mixing by earthworms. Overall, results showed redistribution of tracer ¹⁵N through trees and surface soils without any losses, as whole‐ecosystem recovery remained constant between 1 and 5–6 years at 70% of the ¹⁵N addition. These results demonstrate the persistent ecosystem retention of N deposition even as it redistributes, without additional plant uptake over this timescale.