Nitrogen uptake strategies of edaphically specialized Bornean tree species

The association of tree species with particular soil types contributes to high β diversity in forests, but the mechanisms producing such distributions are still debated. Soil nitrogen (N) often limits growth and occurs in differentially available chemical forms. In a Bornean forest where tree species composition changes dramatically along a soil gradient varying in supplies of different N-forms, we investigated whether tree species’ N-uptake and soil specialization strategies covaried. We analyzed foliar ¹⁵N natural abundance for a total of 216 tree species on clay or sandy loam (the soils at the gradient’s extremes) and conducted a ¹⁵N-tracer experiment with nine specialist and generalist species to test whether species displayed flexible or differential uptake of ammonium and nitrate. Despite variation in ammonium and nitrate supplies and nearly 4 ‰ difference in foliar δ¹⁵N between most soil specialists and populations of generalists on these soils, our ¹⁵N tracer experiment showed little support for the hypothesis that soil specialists vary in N-form use or the ratios in which they use these forms. Instead, our results indicate that these species possess flexible capacities to take up different inorganic N forms. Variation between soil specialists in uptake of different N forms is thus unlikely to cause the soil associations of tree species and high β diversity characteristic of this Bornean rain forest. Flexible uptake strategies would facilitate N-acquisition when supply rates of N-forms exhibit spatiotemporal variation and suggest that these species may be functionally redundant in their responses to N gradients and influences on ecosystem N-cycles.     

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

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

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.     

Distribution and cycling of C, N, Ca, Mg, K and P in three pristine, old-growth forests in the Cordillera de Piuchué, Chile

We assessed a number of biomass and soil parameters in order to examinerelationships among nutrient availability, forest productivity and vegetationpatterns in two old-growth forested watersheds in a pristine montane landscapeon Isla de Chiloé, Chile. We selected watersheds in both gymnosperm- andangiosperm-dominated forests and determined tree species, d.b.h. and health forall trees < 2 cm d.b.h. in plots established at 50m intervals. Soils were sampled at two depths in each plot andanalyzed for total C and N, and for exchangeable Ca, K, Mg andresin-extractableP. Allometric relationships and vegetation nutrient concentrations were used todetermine above-ground pools from the vegetation survey data. Growth rates werederived from increment core measures. Soil pools of most elements measuredappear adequate to support forest growth indefinitely. Mineralized nitrogen,which is similar in quantity to the annual demand for nitrogen from the soil isthe exception, consistent with the possibility of N limitation in two of theforest types studied. A third type, an evergreen broadleaved forest, appears torequire substantially more nitrogen than would appear to be available from netmineralization measurements. Productivity per unit of nitrogen required fromthesoil is quite high, largely as a consequence of the evergreen habit of thespecies in these forests. Compared to other temperate montane forests in theNorthern Hemisphere, nutrient pools and cycling characteristics were found tobemostly similar across forest types, in spite of considerable variation invegetation and soils.     

Organic farm does not improve neither soil,or water quality in rural watersheds from southeastern Brazil

This study was conducted in a rural region where there are conventional and organic farms, the agricultural production includes more than 20 million people, and the effect on environmental quality is still poorly known in terms of indicators. Our objectives were: (1) compare soils attributes to reference areas, (2) verifying if cultivated areas under different farm systems presented differences in the soils attributes, (3) evaluate the attributes of quality water of watersheds and comparing the results with limiting values established by environmental legislation, and (4) analyze the values considering three criterion: watersheds, climatic season, and region of the landscapes. The study was conducted in two rural watersheds that have similar biophysical features and located in the Ibiúna municipality, São Paulo State, Brazil. However, one watershed encompasses farms where landowners largely use conventional agricultural systems. In the other watershed approximately 25% of the farms there are using an organic farm system. In the two watersheds soil samples were collected in sites covered with natural forest and in sites with agriculture (one watershed being organic and other being conventional). The attributes analyzed were soil bulk density (BD), concentrations of Carbon (C) and Nitrogen (N), C:N ratio, C Management Index, and the abundance of ¹³C and ¹⁵N in the soil organic matter. Water attributes were analyzed onsite or in laboratory after analysis of samples. Analyses included: air and water temperature, pH, dissolved oxygen, salinity, total of dissolved solids, total solids, electric conductivity, turbidity, total chloride, nitrate, total phosphorus and potassium. Regarding the soil attributes our database revealed that (1) the soils from cultivated sites of both watersheds presented significant differences from their respective forested areas, (2) Soil attributes are of equal quality in both farm systems. Concerning water attributes: (1) almost all attributes presented values better than the limiting values stipulated by Brazilian legislation; (2) the watersheds did not present significant differences of most of the attributes; (3) in the criteria climatic season data showed some significant differences. The data showed that the soils from the areas used for agricultural ends present belief that significantly worse soil quality in comparison to soils from sites still covered with natural forest. Neither the land cover nor farming system are altering the superficial water quality of the studied watershed and this appears to be related to the extensive percentage of natural remaining vegetation that still exists in both watersheds. The seasonality is an important force that drives the quality characteristics of the water. We highlight that the principles of organic agriculture should be practiced more efficiently and influences such as deforestation should be rigorously avoided.     

Long-term Changes in Forest Carbon and Nitrogen Cycling Caused by an Introduced Pest/Pathogen Complex

Invasion of exotic forest pests and pathogens is a serious environmental problem for many forests throughout the world, and has been especially damaging to forests of eastern North America. We studied the impacts of an exotic pest/pathogen complex, the beech bark disease (BBD), in the Catskill Mountains of New York State, USA. In this region, BBD has caused a decline in the basal area of American beech (Fagus grandifolia Ehrh.) over the last 60 years and this decline has been accompanied by an increase in the basal area of sugar maple (Acer saccharum Marsh.). We studied the impacts of the BBD on carbon (C) and nitrogen (N) cycling using a series of stands that represented a sequence of disease impact and beech replacement by sugar maple. Our study showed that these long-term changes in tree species composition can lead to important changes in C and N cycling in the ecosystem, including an increase in litter decomposition, a decrease in soil C:N ratio, and an increase in extractable nitrate in the soil and nitrate in soil solution. Rates of potential net N mineralization and nitrification did not change across the BBD sequence, but the fraction of mineralized N that was nitrified increased significantly. Many of the observed changes in ecosystem function are larger in magnitude than those attributed to climate change or air pollution, suggesting that the impacts of invasive pests and pathogens on tree species composition could be one of the most important factors driving changes in C and N cycling in these forests in the coming decades.     

Deforestation of watersheds of Panama: nutrient retention and export to streams

A series of eight watersheds on the Pacific coast of Panama where conversion of mature lowland wet forest to pastures by artisanal burning provided watershed-scale experimental units with a wide range of forest cover (23, 29, 47, 56, 66, 73, 73, 91, and 92 %). We used these watersheds as a landscape-scale experiment to assess effects of degree of deforestation on within-watershed retention and hydrological export of atmospheric inputs of nutrients. Retention was estimated by comparing rainfall nutrient concentrations (volume-weighted to allow for evapotranspiration) to concentrations in freshwater reaches of receiving streams. Retention of rain-derived nutrients in these Panama watersheds averaged 77, 85, 80, and 62 % for nitrate, ammonium, dissolved organic N, and phosphate, respectively. Retention of rain-derived inorganic nitrogen, however, depended on watershed cover: retention of nitrate and ammonium in pasture-dominated watersheds was 95 and 98 %, while fully forested watersheds retained 65 and 80 % of atmospheric nitrate and ammonium inputs. Watershed forest cover did not affect retention of dissolved organic nitrogen and phosphate. Exports from more forested watersheds yielded DIN/P near 16, while pasture-dominated watersheds exported N/P near 2. The differences in magnitude of exports and ratios suggest that deforestation in these Panamanian forests results in exports that affect growth of plants and algae in the receiving stream and estuarine ecosystems. Watershed retention of dissolved inorganic nitrogen calculated from wet plus dry atmospheric deposition varied from 90 % in pasture- to 65 % in forest-dominated watersheds, respectively. Discharges of DIN to receiving waters from the watersheds therefore rose from 10 % of atmospheric inputs for pasture-dominated watersheds, to about 35 % of atmospheric inputs for fully forested watersheds. These results from watersheds with no agriculture or urbanization, but different conversion of forest to pasture by burning, show significant, deforestation-dependent retention within tropical watersheds, but also ecologically significant, and deforestation-dependent, exports that are biologically significant because of the paucity of nutrients in receiving tropical stream and coastal waters.     

Nitrogen Addition Increases Carbon Storage in Soils, But Not in Trees, in an Eastern U.S. Deciduous Forest

Forest ecosystems in most industrialized and agricultural regions receive elevated rates of atmospheric nitrogen (N) deposition from air pollution. To evaluate the effects of excess N deposition on carbon (C) and N cycling, we experimentally added N (as NH₄NO₃) to naturally-occurring, single-species plots of five different tree species that are common in the Northern Hardwood forests of northeastern North America: sugar maple (Acer saccharum Marsh), American beech (Fagus grandifolia Ehrh.), yellow birch (Betula alleghaniensis Britton), eastern hemlock (Tsuga canadensis (L.) Carr), and northern red oak (Quercus rubra L.). The experiment was performed in the Catskill Mountains of southeastern New York State, USA, and used a paired-plot design with six replicate plots per species. After 6 years of treatment, most species showed increases in foliar N concentrations in N-treated plots, but only for maple and birch were those increases statistically significant. No significant effects of the N treatment were observed on woody biomass increment or aboveground net primary production (ANPP) for any species. In the oak plots, the N treatment increased acorn production in mast years. In the soils, the N treatment was associated with a significant decline in potential N mineralization and nitrification rates in the mineral horizon but not in the forest floor, and in the mineral horizon the effect of the N treatment varied among species. The N treatment caused a significant increase in C stock, N stock and C:N ratio in the forest floor, with the largest effect in the hemlock plots. Nitrate leaching increased significantly in treated plots compared to controls. Dissolved organic carbon (DOC) in soil solution was unaffected by the N treatment, but the variation in DOC across plots was correlated with the C stock in the forest floor. These results suggest that the ANPP of these forests is not limited by N availability, but that excess N may cause accumulations of C in the forest floor, particularly in hemlock stands, perhaps through inhibition of decomposition rates or by altering phenolic chemistry of the litter. The magnitude, and sometimes the direction of the N treatment responses varied among species, suggesting that predictions of forest responses to elevated N deposition should take into account spatial and temporal variation in tree species composition.     

Nitrogen stable isotopes indicate differences in nitrogen cycling between two contrasting Jamaican montane forests

Background and aims

The aim of this study is to enhance our knowledge of nitrogen (N) cycling and N acquisition in tropical montane forests through analysis of stable N isotopes (δ¹⁵N).

Methods

Leaves from eight common tree species, leaf litter, soils from three depths and roots were sampled from two contrasting montane forest types in Jamaica (mull ridge and mor ridge) and were analysed for δ¹⁵N.

Results

All foliar δ¹⁵N values were negative and varied among the tree species but were significantly more negative in the mor ridge forest (by about 2 ‰). δ¹⁵N of soils and roots were also more negative in mor ridge forests by about 3 ‰. Foliar δ¹⁵N values were closer to that of soil ammonium than soil nitrate suggesting that trees in these forests may have a preference for ammonium; this may explain the high losses of nitrate from similar tropical montane forests. There was no correlation between the rankings of foliar δ¹⁵N in the two forest types suggesting a changing uptake ratio of different N forms between forest types.

Conclusions

These results indicate that N is found at low concentrations in this ecosystem and that there is a tighter N cycle in the mor ridge forest, confirmed by reduced nitrogen availability and lower rates of nitrification. Overall, soil or root δ¹⁵N values are more useful in assessing ecosystem N cycling patterns as different tree species showed differences in foliar δ¹⁵N between the two forest types.     

A Decade of Streamwater Nitrogen and Forest Dynamics after a Mountain Pine Beetle Outbreak at the Fraser Experimental Forest,Colorado

Forests of western North America are currently experiencing extensive tree mortality from a variety of bark beetle species, and insect outbreaks are projected to increase under warmer, drier climates. Unlike the abrupt biogeochemical changes typical after wildfire and timber harvesting, the outcomes of insect outbreaks are poorly understood. The mountain pine bark beetle (Dendroctonus ponderosae) began to attack lodgepole pine (Pinus contorta) at the Fraser Experimental Forest in 2002 and spread throughout the research area by 2007. We compared streamwater nitrogen (N) from 2003 through 2012 with data from the previous two decades in four watersheds with distinct forest management histories, stand structures, and responses to the beetle outbreak. Watersheds dominated by old-growth had larger trees and lost 85% of overstory pine and 44% of total basal area to bark beetles. In contrast, managed watersheds containing a mixture of second-growth (30–60 year old) and old-growth (250- to 350-year old) had higher density of subcanopy trees, smaller mean tree diameter, and lower bark beetle-induced mortality (~26% of total basal area). Streamwater nitrate concentrations were significantly higher in old-growth watersheds during the outbreak than pre-outbreak levels during snowmelt and base flow seasons. In mixed-age stands, streamwater nitrate concentrations were unaffected by the outbreak. Beetle outbreak elevated inorganic N export 43 and 74% in two old-growth watersheds though the amounts of N released in streamwater were low (0.04 and 0.15 kg N ha^?1) relative to atmospheric inputs (<2% of annual N deposition). Increased height, diameter, and foliar N of measured in residual live trees augmented demand for N, far in excess of the change in N export during the outbreak. Reallocation of soil resources released after pine mortality to overstory and understory vegetation helps explain high nutrient retention in watersheds affected by bark beetle outbreaks.     

Nitrogen (N) Dynamics in the Mineral Soil of a Central Appalachian Hardwood Forest During a Quarter Century of Whole-Watershed N Additions

The structure and function of terrestrial ecosystems are maintained by processes that vary with temporal and spatial scale. This study examined temporal and spatial patterns of net nitrogen (N) mineralization and nitrification in mineral soil of three watersheds at the Fernow Experimental Forest, WV: 2 untreated watersheds and 1 watershed receiving aerial applications of N over a 25-year period. Soil was sampled to 5 cm from each of seven plots per watershed and placed in two polyethylene bags—one bag brought to the laboratory for extraction/analysis, and the other bag incubated in situ at a 5 cm depth monthly during growing seasons of 1993–1995, 2002, 2005, 2007–2014. Spatial patterns of net N mineralization and nitrification changed in all watersheds, but were especially evident in the treated watershed, with spatial variability changing non-monotonically, increasing then decreasing markedly. These results support a prediction of the N homogeneity hypothesis that increasing N loads will increase spatial homogeneity in N processing. Temporal patterns for net N mineralization and nitrification were similar for all watersheds, with rates increasing about 25–30% from 1993 to 1995, decreasing by more than 50% by 2005, and then increasing significantly to 2014. The best predictor of these synchronous temporal patterns across all watersheds was number of degree days below 19°C, a value similar to published temperature maxima for net rates of N mineralization and nitrification for these soils. The lack of persistent, detectable differences in net nitrification between watersheds is surprising because fertilization has maintained higher stream-water nitrate concentrations than in the reference watersheds. Lack of differences in net nitrification among watersheds suggests that N-enhanced stream-water nitrate following N fertilization may be the result of a reduced biotic demand for nitrate following fertilization with ammonium sulfate.     

Ecosystem processes and nitrogen export in northern U.S. watersheds

There is much interest in the relationship of atmospheric nitrogen (N) inputs to ecosystem outputs as an indicator of possible "nitrogen saturation" by human activity. Longer-term, ecosystem-level mass balance studies suggest that the relationship is not clear and that other ecosystem processes may dominate variation in N outputs. We have been studying small, forested watershed ecosystems in five northern watersheds for periods up to 35 years. Here I summarize the research on ecosystem processes and the N budget. During the past 2 decades, average wet-precipitation N inputs ranged from about 0.1 to 6 kg N ha(-1) year(-1) among sites. In general, sites with the lowest N inputs had the highest output-to-input ratios. In the Alaska watersheds, streamwater N output exceeded inputs by 70 to 250%. The ratio of mean monthly headwater nitrate (NO3-) concentration to precipitation NO3- concentration declined with increased precipitation concentration. A series of ecosystem processes have been studied and related to N outputs. The most important appear to be seasonal change in hydrologic flowpath, soil freezing, seasonal forest-floor inorganic N pools resulting from over-winter mineralization beneath the snowpack, spatial variation in watershed forest-floor inorganic N pools, the degree to which snowmelt percolates soils, and gross soil N mineralization rates.     

Context-dependent tree species effects on soil nitrogen transformations and related microbial functional genes

Although it is generally accepted that tree species can influence nutrient cycling processes in soils, effects are not consistently found, nor are the mechanisms behind tree species effects well understood. Our objectives were to gain insights into the mechanism(s) underlying the effects of tree species on soil nitrogen cycling processes, and to determine the consistency of tree species effects across sites. We compared N cycling in soils beneath six tree species (ash, sycamore maple, lime, beech, pedunculate oak, Norway spruce) in common garden experiments planted 42 years earlier at three sites in Denmark with distinct land-use histories (forest and agriculture). We measured: (1) net and gross rates of N transformations using the ¹⁵N isotope pool-dilution method, (2) soil microbial community composition through qPCR of fungal ITS, bacterial and archaeal 16S, and (3) abundance of functional genes associated with N cycling processes—for nitrification the archaeal and bacterial ammonia-monooxygenase genes (amoA AOA and amoA AOB, respectively) and for denitrification, the nitrate reductase genes nirK and nirS. Carbon concentrations were higher in soils under spruce than under broadleaves, so N transformation rates were standardized per g soil C. Soil NH₄⁺ parameters (gross ammonification, gross NH₄⁺ consumption, net ammonification (net immobilization in this case), and NH₄⁺ concentrations, per g C) were all lowest in soils under spruce. Soils under spruce also had the lowest gene abundance of bacteria, bacterial:fungal ratio, denitrifying microorganisms, ammonia-oxidizing archaea and ammonia-oxidizing bacteria. Differences in N-cycling processes and organisms among the five broadleaf species were smaller. The ‘spruce effect’ on soil microbes and N transformations appeared to be driven by its acidifying effect on soil and tighter N cycling, which occurred at the previously forested sites but not at the previously agricultural site. We conclude that existing characteristics of soils, including those resulting from previous land use, mediate the effects of tree species on the soil microbial communities and activities that determine rates of N-cycling processes.     

Modeling of nitrogen dynamics in an Austrian alpine forest ecosystem on calcareous soils: a scenario-based risk assessment under changing environmental conditions

We modeled the behavior of an Austrian alpine forest ecosystem on calcareous soils under changing climate and atmospheric nitrogen deposition scenarios. The change of nitrate leaching, emission rates of nitrogen compounds, and forest productivity were calculated using four process-oriented models for the periods 1998-2002 and 2048-2052. Each model reflects with high detail a segment of the ecosystem: PnET-N-DNDC (photosynthesis-evapotranspiration-nitrification-denitrification-decomposition; short-term nitrogen cycling), BROOK90 (water balance for small and homogenous forest watersheds), HYDRUS (water flux in complex and heterogenous soils), and PICUS v1.3 (forest productivity). The nitrogen balance model (NBM) combines the individual results into a comprehensive picture and extends the specific values beyond the limits of the individual models. The evaluation of the findings was outlined with TRACE, a model enabling a long-term prognosis of nitrogen cycling in annual time steps. Temperature increase and nitrogen input are influenced by various components and processes of the forest ecosystem. An increase of the temperature of 2.5 degrees C led to an enhancement of the N2O emission rates and affected the mineralization and the nitrification rates with the consequence of increased nitrate leaching into the subsoil. Enhanced nitrogen input also showed notable effects on nitrate leaching.     

Evidence for a Regime Shift in Nitrogen Export from a Forested Watershed

In this study, we document a functional regime shift in stream inorganic nitrogen (N) processing indicated by a major change in N export from a forested watershed. Evidence from 36 years of data following experimental clearcut logging at Coweeta Hydrologic Laboratory, NC, suggests that forest disturbance in this area can cause elevation of dissolved inorganic N (DIN) loss lasting decades or perhaps longer. This elevation of N export was apparently caused by an initial pulse of organic matter input, reduced vegetation uptake, increased mineralization of soil organic N, and N fixation by black locust-associated bacteria following clearcut logging. In forested reference watersheds at Coweeta, maximum DIN concentration occurs in summer when base flow is low, but the clearcut watershed shifted to a pattern of maximum winter DIN concentration. The seasonal pattern of DIN concentration and export from reference watersheds can be explained by terrestrial and in-stream processes, but following clearcutting, elevated DIN availability saturated both terrestrial and in-stream uptake, and the N export regime became dominated by hydrologic transport. We suggest that the long-term elevation of stream DIN concentration and export along with the changes in seasonality of DIN export and the relationship between concentration and discharge represent a functional regime shift initiated by forest disturbance.     

A New Conceptual Model of Nitrogen Saturation Based on Experimental Nitrogen Addition to an Oak Forest

The dominant conceptual model of nitrogen (N) saturation in forests predicts the temporal patterns of key N cycling indicators as an initially N-limited forest is progressively enriched in N. We present the results from a long-term N addition experiment in an oak forest in southeastern New York State, USA, which do not conform to the predictions of the conceptual model in several ways. In contrast to the predictions of the conceptual model, the foliar N concentrations in the N-treated stands of our study increased to about 20% above the levels in the control stands and then remained essentially constant, and nitrogen leaching from the treated stands increased almost immediately after the start of the experiment, prior to the onset of elevated nitrification. Concentrations of N in soil solution of the N-treated stands peaked at over 150-fold greater than the concentrations in the control stands. There were no significant changes in potential net N mineralization. Tree mortality increased in the treated stands, but the tree mortality did not appear to be the primary cause of the excess nitrate leaching. Based on these results and those of other recent studies, we present a new conceptual model of the N saturation process focused on the mass balance of N rather than the temporal dynamics of N cycling indicators. The mass balance is characterized by inputs of N from atmospheric deposition and fertilization, internal sinks in the vegetation and soils, and outputs to leaching and gaseous losses. The key points of the conceptual model are (1) added N can flow simultaneously to all sinks and losses in the system, (2) the fate of the added N and the temporal patterns of flow of N depend on the strength of the sinks and the factors that control them, and (3) the movement of N to the various sinks determines how N saturation is manifested in the ecosystem. We distinguish capacity N saturation, in which the sinks in the vegetation and soil are zero or negative, from kinetic N saturation, in which the sinks are positive but lower than the N input rate. The sink strengths in the vegetation and soil have two components, one due to carbon (C) accumulation in the system and the other due to change in the stoichiometry (C:N ratio) of the pool. Further work quantifying the magnitudes and controlling factors for the N sinks will allow better prediction of the dynamics of N saturation in different types of forested ecosystems.     

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.     

The effect of single tree species on soil microbial activities related to C and N cycling in the Siberian artificial afforestation experiment

The effects of grassland conversion to forest vegetation and of individual tree species on microbial activity in Siberia are largely unstudied. Here, we examined the effects of the six most commonly dominant tree species in Siberian forests (Scots pine, spruce, Arolla pine, larch, aspen and birch) on soil C and N mineralization, N₂O-reduction and N₂O production during denitrification 30 years after planting. We also documented the effect of grassland conversion to different tree species on microbial activities at different soil depths and their relationships to soil chemical properties. The effects of tree species and grassland conversion were more pronounced on N than on C transformations. Tree species and grassland conversion did significantly alter substrate-induced respiration (SIR) and basal respiration, but the differences were not as large as those observed for N transformations. Variances in SIR and basal respiration within species were markedly lower than those in N transformations. Net N mineralization, net nitrification, and denitrification potential were highest under Arolla pine and larch, intermediate under deciduous aspen and birch, and lowest beneath spruce and Scots pine. Tree species caused similar effects on denitrification potential, net N mineralization, and net nitrification, but effects on N₂O reduction rate were idiosyncratic, indicating a decoupling of N₂O production and reduction. We predict that deciduous species should produce more N₂O in the field than conifers, and that Siberian forests will produce more N₂O if global climate change alters tree species composition. Basal respiration and SIR showed inverse responses to tree species: when basal respiration increased in response to a given tree species, SIR declined. SIR may have been controlled by NH₄ ⁺ availability and related therefore to N mineralization, which was negatively affected by grassland conversion. Basal respiration appeared to be less limited by NH₄ ⁺ and controlled mostly by readily available organic C (DOC), which was higher in concentration under forests than in grassland and therefore basal respiration was higher in forested soils. We conclude that in the Siberian artificial afforestation experiment, soil C mineralization was not limited by N.     

Seasonal changes in the chemical quality and biodegradability of dissolved organic matter exported from soils to streams in coastal temperate rainforest watersheds

The composition and biodegradability of streamwater dissolved organic matter (DOM) varies with source material and degree of transformation. We combined PARAFAC modeling of fluorescence excitation–emission spectroscopy and biodegradable dissolved organic carbon (BDOC) incubations to investigate seasonal changes in the lability of DOM along a soil-stream continuum in three soil types: bog, forested wetland and upland forest. The percent BDOC ranged from 7 to 38% across all sites, and was significantly greater in soil compared to streamwater in the bog and forested wetland, but not in the upland forest. The percent BDOC also varied significantly over the entire sampling period in soil and streamwater for the bog and forested wetland, as BDOC peaked during the spring runoff and was lowest during the summer months. Moreover, the chemical quality of DOM in wetland soil and streamwater was similar during the spring runoff and fall wet season, as demonstrated by the similar contribution of protein-like fluorescence (sum of tyrosine and tryptophan fluorescence) in soil water and in streams. These findings suggest that the tight coupling between terrestrial and aquatic ecosystems is responsible for the delivery of labile DOM from wetland soils to streams. The contribution of protein-like fluorescence was significantly correlated with BDOC (p < 0.001) over the entire sampling period indicating DOM is an important source of C and N for heterotrophic microbes. Taken together, our findings suggest that the production of protein-rich, labile DOM and subsequent loss in stream runoff might be an important loss of labile C and N from coastal temperate watersheds.     

Nitrogen deposition and leaching from two forested catchments in Southwest China--preliminary data and research needs

Increased nitrogen deposition has resulted in increased nitrogen pools and nitrogen leaching in European and North American forest soils. The development in Asia in general, and China in particular, suggests increased deposition of reduced nitrogen from changes in agricultural practices and of oxidized nitrogen from rapid growth of the transportation sector. Decreased nitrogen retention in forested areas in the future may cause increased NO3- leaching and, thus, acidification and eutrophication in surface waters. The differences in climate, ecosystems, land use, and deposition history make direct application of knowledge from studies in Europe and North America difficult. In Southwest China the potential for nitrogen mobilization from forest soils may be high because of the warm and humid climate, resulting in high decomposition rates of soil organic matter. However, there are very few data available for quantifying the suspected potential for increased nitrogen leaching in forest ecosystems. Here we present data from two forested catchments, dominated by Masson pine (Pinus massoniana), near Guiyang and Chongqing, respectively, in Southwest China. The present nitrogen deposition is moderate, estimated in the range from 10 to 40 kg N ha(-1) year(-1). The C/N ratios of the soils are generally below 15. Nitrate concentrations in soil water are rather variable in space, with highest values of several hundred microequivalents per liter. The turnover rate of nitrogen in the forest ecosystem is quite high compared to the atmospheric deposition rate. At present, nitrate runoff from the catchments is low and intermediate in Guiyang and Chongqing, respectively. More research is needed to improve our ability to predict future nitrogen leaching from subtropical Asian coniferous forests.