施氮与凋落物去除影响下中亚热带阔叶林土壤氮素矿化潜势和硝化潜势研究

氮沉降影响土壤氮循环,而凋落物是土壤有机氮的主要来源,因此,为了探讨氮沉降和凋落物是否去除作用下,亚热带森林土壤潜在的氮素矿化与硝化作用,选择已进行8年模拟氮沉降试验的亚热带罗浮栲(Castanopsis fabri)常绿阔叶林土壤为研究对象,野外样地氮添加设置3个水平:对照(CK,0 kg·hm^?2·a^?1)、低氮(LN,75 kg·hm^?2·a^?1)、高氮(HN,150 kg·hm^?2·a^?1),两种凋落物管理方式(保留凋落物,L和去除凋落物,R),土壤采样后,通过室内间歇淋洗好气培养法,研究土壤氮素矿化潜势差异,以及不同底物条件下(铵态氮水平:N 0,100、150、200 mg·kg^?1)土壤硝化潜势的差异。结果表明:土壤氮素快速矿化主要在培养前7 d,矿化累积量(Nt)为102.81—153.71 mg·kg^?1,矿化潜势(N0)范围为193.84—289.80 mg·kg^?1,N0依次为:保留凋落物低氮(LN-L)>保留凋落物对照(CK-L)>去除凋落物低氮(LN-R)>去除凋落物对照(CK-R)>去除凋落物高氮(HN-R)>保留凋落物高氮(HN-L);两种凋落物处理方式下,LN水平土壤的Nt与N0均高于CK、HN。保留凋落物情况下,有较高的土壤硝化潜势;在无添加硝化底物(铵态氮水平为N 0 mg·kg^?1)的条件下,野外氮添加水平高的土壤硝化潜势也高;而在添加不同硝化底物(铵态氮)的条件下,土壤硝化潜势并未随硝化底物水平的增加而增加,且硝化底物水平为N 100 mg·kg^?1时硝化潜势最大。研究表明,虽然保留凋落物可以增加土壤氮矿化潜势,而氮沉降则影响氮矿化潜势。当研究土壤硝化潜势时,应当根据土壤类型等因素选择合适的硝化底物(铵态氮)添加量。     

Dynamics of nitrogen and dissolved organic carbon at the Hubbard brook experimental forest

The factors controlling spatial and temporal patterns in soil solution and streamwater chemistry are highly uncertain in northern hardwood forest ecosystems in the northeastern United States, where concentrations of reactive nitrogen (Nr) in streams have surprisingly declined over recent decades in the face of persistent high rates of atmospheric Nr deposition and aging forests. Reactive nitrogen includes inorganic species (e.g., ammonium [NH4+], nitrate [NO3-]) and some organic forms (e.g., amino acids) available to support the growth of plants and microbes. The objective of this study was to examine controls on the spatial and temporal patterns in the concentrations and fluxes of nitrogen (N) species and dissolved organic carbon (DOC) in a 12-year record of soil solutions and streamwater along an elevational gradient (540-800 m) of a forested watershed at the Hubbard Brook Experimental Forest (HBEF) in the White Mountains of New Hampshire, USA. Dissolved organic N and DOC concentrations were elevated in the high-elevation spruce-fir-white birch (SFB) zone of the watershed, while NO3- was the dominant N species in the lower elevation hardwood portion of the watershed. Within the soil profile, N retention was centered in the mineral horizon, and significant amounts of N were retained between the lower mineral soil and the stream, supporting the idea that near- and in-stream processes are significant sinks for N at the HBEF. Temporal analysis suggested that hydrologic flow paths can override both abiotic and biotic retention mechanisms (i.e., during the non-growing season when most hydrologic export occurs, or during years with high rainfall), there appears to be direct flushing of N from the organic horizons into the stream via horizontal flow. Significant correlations between soil NO3- concentrations, nitrification rates and streamwater NO3- exports show the importance of biological production as a regulator of inorganic N export. The lack of internal production response (e.g., mineralization, nitrification) to a severe ice storm in 1998 reinforces the idea that plant uptake is the dominant regulator of export response to disturbance.     

Microbial nitrogen limitation increases decomposition

With anthropogenic nutrient inputs to ecosystems increasing globally, there are long-standing, fundamental questions about the role of nutrients in the decomposition of organic matter. We tested the effects of exogenous nitrogen and phosphorus inputs on litter decomposition across a broad suite of litter and soil types. In one experiment, C mineralization was compared across a wide array of plants individually added to a single soil, while in the second, C mineralization from a single substrate was compared across 50 soils. Counter to basic stoichiometric decomposition theory, low N availability can increase litter decomposition as microbes use labile substrates to acquire N from recalcitrant organic matter. This "microbial nitrogen mining" is consistently suppressed by high soil N supply or substrate N concentrations. There is no evidence for phosphorus mining as P fertilization increases short- and long-term mineralization. These results suggest that basic stoichiometric decomposition theory needs to be revised and ecosystem models restructured accordingly in order to predict ecosystem carbon storage responses to anthropogenic changes in nutrient availability.     

Stoichiometric controls of nitrogen and phosphorus cycling in decomposing beech leaf litter

Resource stoichiometry (C:N:P) is an important determinant of litter decomposition. However, the effect of elemental stoichiometry on the gross rates of microbial N and P cycling processes during litter decomposition is unknown. In a mesocosm experiment, beech (Fagus sylvatica L.) litter with natural differences in elemental stoichiometry (C:N:P) was incubated under constant environmental conditions. After three and six months, we measured various aspects of nitrogen and phosphorus cycling. We found that gross protein depolymerization, N mineralization (ammonification), and nitrification rates were negatively related to litter C:N. Rates of P mineralization were negatively correlated with litter C:P. The negative correlations with litter C:N were stronger for inorganic N cycling processes than for gross protein depolymerization, indicating that the effect of resource stoichiometry on intracellular processes was stronger than on processes catalyzed by extracellular enzymes. Consistent with this, extracellular protein depolymerization was mainly limited by substrate availability and less so by the amount of protease. Strong positive correlations between the interconnected N and P pools and the respective production and consumption processes pointed to feed-forward control of microbial litter N and P cycling. A negative relationship between litter C:N and phosphatase activity (and between litter C:P and protease activity) demonstrated that microbes tended to allocate carbon and nutrients in ample supply into the production of extracellular enzymes to mine for the nutrient that is more limiting. Overall, the study demonstrated a strong effect of litter stoichiometry (C:N:P) on gross processes of microbial N and P cycling in decomposing litter; mineralization of N and P were tightly coupled to assist in maintaining cellular homeostasis of litter microbial communities.     

Nitrogen dynamics in an Australian semiarid grassland soil

We conducted a four-week laboratory incubation of soil from a Themeda triandra Forsskal grassland to clarify mechanisms of nitrogen (N) cycling processes in relation to carbon (C) and N availability in a hot, semiarid environment. Variation in soil C and N availability was achieved by collecting soil from either under tussocks or the bare soil between tussocks, and by amending soil with Themeda litter. We measured N cycling by monitoring: dissolved organic nitrogen (DON), ammonium (NH4+), and nitrate (NO3-) contents, gross rates of N mineralization and microbial re-mineralization, NH4+ and NO3- immobilization, and autotrophic and heterotrophic nitrification. We monitored C availability by measuring cumulative soil respiration and dissolved organic C (DOC). Litter-amended soil had cumulative respiration that was eightfold greater than non-amended soil (2000 compared with 250 microg C/g soil) and almost twice the DOC content (54 compared with 28 microg C/g soil). However, litter-amended soils had only half as much DON accumulation as non-amended soils (9 compared with 17 microg N/g soil) and lower gross N rates (1-4 compared with 13-26 microg N x [g soil](-1) x d(-1)) and NO3- accumulation (0.5 compared with 22 microg N/g soil). Unamended soil from under tussocks had almost twice the soil respiration as soil from between tussocks (300 compared with 175 microg C/g soil), and greater DOC content (33 compared with 24 microg C/g soil). However, unamended soil from under tussocks had lower gross N rates (3-20 compared with 17-31 microg N x [g soil](-1) d(-1)) and NO3- accumulation (18 compared with 25 microg N/g soil) relative to soil from between tussocks. We conclude that N cycling in this grassland is mediated by both C and N limitations that arise from the patchiness of tussocks and seasonal variability in Themeda litterfall. Heterotrophic nitrification rate explained >50% of total nitrification, but this percentage was not affected by proximity to tussocks or litter amendment. A conceptual model that considers DON as central to N cycling processes provided a useful initial framework to explain results of our study. However, to fully explain N cycling in this semiarid grassland soil, the production of NO3- from organic N sources must be included in this model.     

Potential nitrogen constraints on soil carbon sequestration under low and elevated atmospheric CO2

The interaction between nitrogen cycling and carbon sequestration is critical in predicting the consequences of anthropogenic increases in atmospheric CO2 (hereafter, Ca). The progressive N limitation (PNL) theory predicts that carbon sequestration in plants and soils with rising Ca may be constrained by the availability of nitrogen in many ecosystems. Here we report on the interaction between C and N dynamics during a four-year field experiment in which an intact C3/C4 grassland was exposed to a gradient in Ca from 200 to 560 micromol/mol. There were strong species effects on decomposition dynamics, with C loss positively correlated and N mineralization negatively correlated with Ca for litter of the C3 forb Solanum dimidiatum, whereas decomposition of litter from the C4 grass Bothriochloa ischaemum was unresponsive to Ca. Both soil microbial biomass and soil respiration rates exhibited a nonlinear response to Ca, reaching a maximum at approximately 440 micromol/mol Ca. We found a general movement of N out of soil organic matter and into aboveground plant biomass with increased Ca. Within soils we found evidence of C loss from recalcitrant soil C fractions with narrow C:N ratios to more labile soil fractions with broader C:N ratios, potentially due to decreases in N availability. The observed reallocation of N from soil to plants over the last three years of the experiment supports the PNL theory that reductions in N availability with rising Ca could initially be overcome by a transfer of N from low C:N ratio fractions to those with higher C:N ratios. Although the transfer of N allowed plant production to increase with increasing Ca, there was no net soil C sequestration at elevated Ca, presumably because relatively stable C is being decomposed to meet microbial and plant N requirements. Ultimately, if the C gained by increased plant production is rapidly lost through decomposition, the shift in N from older soil organic matter to rapidly decomposing plant tissue may limit net C sequestration with increased plant production.     

Nitrogen critical loads for alpine vegetation and terrestrial ecosystem response: are we there yet?

Increases in the deposition of anthropogenic nitrogen (N) have been linked to several terrestrial ecological changes, including soil biogeochemistry, plant stress susceptibility, and community diversity. Recognizing the need to identify sensitive indicators of biotic response to N deposition, we empirically estimated the N critical load for changes in alpine plant community composition and compared this with the estimated critical load for soil indicators of ecological change. We also measured the degree to which alpine vegetation may serve as a sink for anthropogenic N and how much plant sequestration is related to changes in species composition. We addressed these research goals by adding 20, 40, or 60 kg N x ha(-1) x yr(-1), along with an ambient control (6 kg N x ha(-1) x yr(-1) total deposition), to a species-rich alpine dry meadow for an eight-year period. Change in plant species composition associated with the treatments occurred within three years of the initiation of the experiment and were significant at all levels of N addition. Using individual species abundance changes and ordination scores, we estimated the N critical loads (total deposition) for (1) change in individual species to be 4 kg N x ha(-1) yr(-1) and (2) for overall community change to be 10 kg N x ha(-1) x yr(-1). In contrast, increases in NO3- leaching, soil solution inorganic NO3-, and net N nitrification occurred at levels above 20 kg N x ha(-1) x yr(-1). Increases in total aboveground biomass were modest and transient, occurring in only one of the three years measured. Vegetative uptake of N increased significantly, primarily as a result of increasing tissue N concentrations and biomass increases in subdominant species. Aboveground vegetative uptake of N accounted for <40% of the N added. The results of this experiment indicate that changes in vegetation composition will precede detectable changes in more traditionally used soil indicators of ecosystem responses to N deposition and that changes in species composition are probably ongoing in alpine dry meadows of the Front Range of the Colorado Rocky Mountains. Feedbacks to soil N cycling associated with changes in litter quality and species composition may result in only short-term increases in vegetation N pools.     

Bromus tectorum invasion alters nitrogen dynamics in an undisturbed arid grassland ecosystem

The nonnative annual grass Bromus tectorum has successfully replaced native vegetation in many arid and semiarid ecosystems. Initial introductions accompanied grazing and agriculture, making it difficult to separate the effects of invasion from physical disturbance. This study examined N dynamics in two recently invaded, undisturbed vegetation associations (C3 and C4). The response of these communities was compared to an invaded/ disturbed grassland. The invaded/disturbed communities had higher surface NH4+ input in spring, whereas there were no differences for surface input of NO3-. Soil inorganic N was dominated by NH4+, but invaded sites had greater subsurface soil NO3-. Invaded sites had greater total soil N at the surface four years post-invasion in undisturbed communities, but total N was lower in the invaded/disturbed communities. Soil delta15N increased with depth in the noninvaded and recently invaded communities, whereas the invaded/disturbed communities exhibited the opposite pattern. Enriched foliar delta15N values suggest that Bromus assimilated subsurface NO3-, whereas the native grasses were restricted to surface N. A Rayleigh distillation model accurately described decomposition patterns in the noninvaded communities where soil N loss is accompanied by increasing soil delta15N; however, the invaded/ disturbed communities exhibited the opposite pattern, suggesting redistribution of N within the soil profile. This study suggests that invasion has altered the mechanisms driving nitrogen dynamics. Bromus litter decomposition and soil NO3- concentrations were greater in the invaded communities during periods of ample precipitation, and NO3- leached from the surface litter, where it was assimilated by Bromus. The primary source of N input in these communities is a biological soil crust that is removed with disturbance, and the lack of N input by the biological soil crust did not balance N loss, resulting in reduced total N in the invaded/disturbed communities. Bromus produced a positive feedback loop by leaching NO3- from decomposing Bromus litter to subsurface soil layers, accessing that deepsoil N pool with deep roots and returning that N to the surface as biomass and subsequent litter. Lack of new inputs combined with continued loss will result in lower total soil N, evidenced by the lower total soil N in the invaded/disturbed communities.     

Seasonal variations in plant species effects on soil N and P dynamics

It is well established that plant species influence ecosystem processes, but we have little ability to predict which vegetation changes will alter ecosystems, or how the effects of a given species might vary seasonally. We established monocultures of eight plant species in a California grassland in order to determine the plant traits that account for species impacts on nitrogen and phosphorus cycling. Plant species differed in their effects on net N mineralization and nitrification rates, and the patterns of species differences varied seasonally. Soil PO4- and microbial P were more strongly affected by slope position than by species. Although most studies focus on litter chemistry as the main determinant of plant species effects on nutrient cycling, this study showed that plant species affected biogeochemical cycling through many traits, including direct traits (litter chemistry and biomass, live-tissue chemistry and biomass) and indirect traits (plant modification of soil bioavailable C and soil microclimate). In fact, species significantly altered N and P cycling even without litter inputs. It became particularly critical to consider the effects of these multiple traits in order to account for seasonal changes in plant species effects on ecosystems. For example, species effects on potential rates of net N mineralization were most strongly influenced by soil bioavailable C in the fall and by litter chemistry in the winter and spring. Under field conditions, species effects on soil microclimate influenced rates of mineralization and nitrification, with species effects on soil temperature being critical in the fall and species effects on soil moisture being important in the dry spring. Overall, this study clearly demonstrated that in order to gain a mechanistic, predictive understanding of plant species effects on ecosystems, it is critical to look beyond plant litter chemistry and to incorporate the effects of multiple plant traits on ecosystems.     

Litter decomposition and nitrogen mineralization from an annual to a monthly model

The forest litter decomposition model (FLDM) described in this paper provides an important basis for assessing the impacts of forest management on seasonal stream water quality and export of dissolved organic carbon (DOC). By definition, models with annual time steps are unable to capture seasonal, within-year variation. In order to simulate seasonal variation in litter decomposition and DOC production and export, we have modified an existing annual FLDM to account for monthly dynamics of decomposition and residual mass in experimental litterbags placed in 21 different forests across Canada.The original annual FLDM was formulated with three main litter pools (fast, slow, and very slow decomposing litter) to address the fact that forest litter is naturally composed of a mixture of organic compounds that decompose at different rates. The annual FLDM was shown to provide better simulations than more complex models like CENTURY and SOMM.The revised monthly model retains the original structure of the annual FLDM, but separates litter decomposition from nitrogen (N) mineralization. In the model, monthly soil temperature, soil moisture, and mean January soil temperature are shown to be the most important controlling variables of within-year variation in decomposition. Use of the three variables in a process-based definition of litter decomposition is a significant departure from the empirical definition in the annual model. The revised model is shown to give similar calculations of residual mass and N concentration as the annual model (r² = 0.91, 0.78), despite producing very different timeseries of decomposition over six years. It is shown from a modelling perspective that (i) forest litter decomposition is independent of N mineralization, whereas N mineralization is dependent on litter decomposition, and (ii) mean January soil temperature defines litter decomposition in the summer because of winter-temperatures’ role in modifying forest-floor microorganism community composition and functioning in the following summer.     

Atmospheric deposition, mineralization and leaching of nitrogen in subtropical forested catchments, South China

In recent years, China has conducted considerable research focusing on the emission and effects of sulphur (S) on human health and ecosystems. By contrast, there has been little emphasis on anthropogenic nitrogen (N) so far, even though studies conducted abroad indicate that long-range atmospheric transport of N and ecological effects (e.g. acidification of soil and water) may be significant. The Sino-Norwegian project IMPACTS, launched in 1999, has established monitoring sites at five forest ecosystems in the southern part of PR China to collect comprehensive data on air quality, acidification status and ecological effects. Here we present initial results about N dynamics at two of the IMPACTS sites located near Chongqing and Changsha, including estimation of atmospheric deposition fluxes of NOx and NHx and soil N transformations. Nitrogen deposition is high at both sites when compared with values from Europe and North America (25-38 kg ha(-1) yr(-1)). About 70% of the deposited N comes as NH4, probably derived from agriculture. Leaching of N from soils is high and nearly all as NO3-. Transformation of N to NO3- in soils results in acidification rates that are high compared to rates found elsewhere. Despite considerable leaching of NO3- from the root zone of the soils, little NO3- appears in streamwater. This indicates that N retention or denitrification, both causing acid neutralization, may be important and probably occur in the groundwater and groundwater discharge zones. The soil flux density of mineral N, which is the sum of N deposition and N mineralization, and which is dominated by the N mineralization flux, may be a good indicator for leaching of NO3- in soils. However, this indicator seems site specific probably due to differences in land-use history and current N requirement.     

A process-based model of nitrogen cycling in forest plantations: Part II. Simulating growth and nitrogen mineralisation of Eucalyptus globulus plantations in south-western Australia

We applied a new version of the G’DAY ecosystem model to short-rotation plantations of Eucalyptus globulus growing under a Mediterranean climate in south-western Australia. The new version, that includes modified submodels for biomass production, water balance, litter and soil organic matter (SOM) decomposition, and soil inorganic N balance, was parameterised and applied to three experimental eucalypt sites (Mumballup, Darkan and Northcliffe) of contrasting productivity. With a common base set of parameter values, the model was able to correctly reproduce observed time series of soil water content, canopy leaf area index and stemwood data at the three sites. The model's ability to simulate soil N supply under forest plantations was tested by simulating N mineralisation at each of the three sites over the duration of the experiment (10 years). Simulated annual net N mineralisation in the litter and top 20 cm soil layer ranged from 50 to 170 kg N ha⁻¹ across the sites as a result of differences in rates of litter production, SOM and litter decomposition, and microbial N immobilisation and (re-)mineralisation. Simulations of annual soil N mineralisation were similar to measured rates over a 3-year period, except for an overestimation in 1 year at Mumballup and 2 years at Darkan. Model results indicated the importance of fine root production and turnover for N supply. As plantations age, supply of N to trees increasingly originates from litter decomposition, while the contribution from decomposition of SOM decreases. Although major soil feedbacks associated with litter production, decomposition and N availability are adequately integrated into G’DAY, further work is required in some aspects of the model, including the utility of the C-allocation submodel over a wide range of site conditions and silvicultural treatments.     

Plants control the seasonal dynamics of microbial N cycling in a beech forest soil by belowground C allocation

Soil microbes in temperate forest ecosystems are able to cycle several hundreds of kilograms of N per hectare per year and are therefore of paramount importance for N retention. Belowground C allocation by trees is an important driver of seasonal microbial dynamics and may thus directly affect N transformation processes over the course of the year. Our study aimed at unraveling plant controls on soil N cycling in a temperate beech forest at a high temporal resolution over a time period of two years, by investigating the effects of tree girdling on microbial N turnover. In both years of the experiment, we discovered (1) a summer N mineralization phase (between July and August) and (2) a winter N immobilization phase (November-February). The summer mineralization phase was characterized by a high N mineralization activity, low microbial N uptake, and a subsequent high N availability in the soil. During the autumn/winter N immobilization phase, gross N mineralization rates were low, and microbial N uptake exceeded microbial N mineralization, which led to high levels of N in the microbial biomass and low N availability in the soil. The observed immobilization phase during the winter may play a crucial role for ecosystem functioning, since it could protect dissolved N that is produced by autumn litter degradation from being lost from the ecosystem during the phase when plants are mostly inactive. The difference between microbial biomass N levels in winter and spring equals 38 kg N/ha and may thus account for almost one-third of the annual plant N demand. Tree girdling strongly affected annual N cycling: the winter N immobilization phase disappeared in girdled plots (microbial N uptake and microbial biomass N were significantly reduced, while the amount of available N in the soil solution was enhanced). This was correlated to a reduced fungal abundance in autumn in girdled plots. By releasing recently fixed photosynthates to the soil, plants may thus actively control the annual microbial N cycle. Tree belowground C allocation increases N accumulation in microorganisms during the winter which may ultimately feed back on plant N availability in the following growing season.     

Tree species effects on decomposition and forest floor dynamics in a common garden

We studied the effects of tree species on leaf litter decomposition and forest floor dynamics in a common garden experiment of 14 tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Betula pendula, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pinus nigra, Pinus sylvestris, Pseudotsuga menziesii, Quercus robur, Quercus rubra, and Tilia cordata) in southwestern Poland. We used three simultaneous litter bag experiments to tease apart species effects on decomposition via leaf litter chemistry vs. effects on the decomposition environment. Decomposition rates of litter in its plot of origin were negatively correlated with litter lignin and positively correlated with mean annual soil temperature (MAT(soil)) across species. Likewise, decomposition of a common litter type across all plots was positively associated with MAT(soil), and decomposition of litter from all plots in a common plot was negatively related to litter lignin but positively related to litter Ca. Taken together, these results indicate that tree species influenced microbial decomposition primarily via differences in litter lignin (and secondarily, via differences in litter Ca), with high-lignin (and low-Ca) species decomposing most slowly, and by affecting MAT(soil), with warmer plots exhibiting more rapid decomposition. In addition to litter bag experiments, we examined forest floor dynamics in each plot by mass balance, since earthworms were a known component of these forest stands and their access to litter in litter bags was limited. Forest floor removal rates estimated from mass balance were positively related to leaf litter Ca (and unrelated to decay rates obtained using litter bags). Litter Ca, in turn, was positively related to the abundance of earthworms, particularly Lumbricus terrestris. Thus, while species influence microbially mediated decomposition primarily through differences in litter lignin, differences among species in litter Ca are most important in determining species effects on forest floor leaf litter dynamics among these 14 tree species, apparently because of the influence of litter Ca on earthworm activity. The overall influence of these tree species on leaf litter decomposition via effects on both microbial and faunal processing will only become clear when we can quantify the decay dynamics of litter that is translocated belowground by earthworms.     

Frequent fire alters nitrogen transformations in ponderosa pine stands of the inland northwest

Recurrent, low-severity fire in ponderosa pine (Pinus ponderosa)/interior Douglas-fir (Pseudotsuga menziesii var. glauca) forests is thought to have directly influenced nitrogen (N) cycling and availability. However, no studies to date have investigated the influence of natural fire intervals on soil processes in undisturbed forests, thereby limiting our ability to understand ecological processes and successional dynamics in this important ecosystem of the Rocky Mountain West. Here, we tested the standing hypothesis that recurrent fire in ponderosa pine/Douglas-fir forests of the Inland Northwest decreases total soil N, but increases N turnover and nutrient availability. We compared soils in stands unburned over the past 69-130 years vs. stands exposed to two or more fires over the last 130 years at seven distinct locations in two wilderness areas. Mineral soil samples were collected from each of the seven sites in June and July of 2003 and analyzed for pH, total C and N, potentially mineralizable N (PMN), and extractable NH4+, NO3-, PO4(-3), Ca+2, Mg+2, and K+. Nitrogen transformations were assessed at five sites by installing ionic resin capsules in the mineral soil in August of 2003 and by conducting laboratory assays of nitrification potential and net nitrification in aerobic incubations. Total N and PMN decreased in stands subjected to multiple fires. This loss of total N and labile N was not reflected in concentrations of extractable NH4+ and NO3-. Rather, multiple fires caused an increase in NO3 sorbed on ionic resins, nitrification potential, and net nitrification in spite of the burned stands not having been exposed to fire for at least 12-17 years. Charcoal collected from a recent fire site and added to unburned soils increased nitrification potential, suggesting that the decrease of charcoal in the absence of fire may play an important role in N transformations in fire-dependent ecosystems in the long term. Interestingly, we found no consistent effect of fire frequency on extractable P or alkaline metal concentrations. Our results corroborate the largely untested hypothesis that frequent fire in ponderosa pine forests increases inorganic N availability in the long term and emphasize the need to study natural, unmanaged sites in far greater detail.     

No evidence that chronic nitrogen additions increase photosynthesis in mature sugar maple forests

Atmospheric nitrogen (N) deposition can increase forest growth. Because N deposition commonly increases foliar N concentrations, it is thought that this increase in forest growth is a consequence of enhanced leaf-level photosynthesis. However, tests of this mechanism have been infrequent, and increases in photosynthesis have not been consistently observed in mature forests subject to chronic N deposition. In four mature northern hardwood forests in the north-central United States, chronic N additions (30 kg N ha(-1) yr(-1) as NaNO3 for 14 years) have increased aboveground growth but have not affected canopy leaf biomass or leaf area index. In order to understand the mechanism behind the increases in growth, we hypothesized that the NO3(-) additions increased foliar N concentrations and leaf-level photosynthesis in the dominant species in these forests (sugar maple, Acer saccharum). The NO3(-) additions significantly increased foliar N. However, there was no significant difference between the ambient and +NO3(-) treatments in two seasons (2006-2007) of instantaneous measurements of photosynthesis from either canopy towers or excised branches. In measurements on excised branches, photosynthetic nitrogen use efficiency (micromol CO2 s(-1) g(-1) N) was significantly decreased (-13%) by NO3(-) additions. Furthermore, we found no consistent NO3(-) effect across all sites in either current foliage or leaf litter collected annually throughout the study (1993-2007) and analyzed for delta 13C and delta 18O, isotopes that can be used together to integrate changes in photosynthesis over time. We observed a small but significant NO3(-) effect on the average area and mass of individual leaves from the excised branches, but these differences varied by site and were countered by changes in leaf number. These photosynthesis and leaf area data together suggest that NO3(-) additions have not stimulated photosynthesis. There is no evidence that nutrient deficiencies have developed at these sites, so unlike other studies of photosynthesis in N-saturated forests, we cannot attribute the lack of a stimulation of photosynthesis to nutrient limitations. Rather than increases in C assimilation, the observed increases in aboveground growth at our study sites are more likely due to shifts in C allocation.     

Biogeochemistry of a temperate forest nitrogen gradient

Wide natural gradients of soil nitrogen (N) can be used to examine fundamental relationships between plant-soil-microbial N cycling and hydrologic N loss, and to test N-saturation theory as a general framework for understanding ecosystem N dynamics. We characterized plant production, N uptake and return in litterfall, soil gross and net N mineralization rates, and hydrologic N losses of nine Douglas-fir (Pseudotsuga menziesii) forests across a wide soil N gradient in the Oregon Coast Range (U.S.A.). Surface mineral soil N (0-10 cm) ranged nearly three-fold from 0.29% to 0.78% N, and in contrast to predictions of N-saturation theory, was linearly related to 10-fold variation in net N mineralization, from 8 to 82 kg N.ha(-1) x yr(-1). Net N mineralization was unrelated to soil C:N, soil texture, precipitation, and temperature differences among sites. Net nitrification was negatively related to soil pH, and accounted for <20% of net N mineralization at low-N sites, increasing to 85-100% of net N mineralization at intermediate- and high-N sites. The ratio of net: gross N mineralization and nitrification increased along the gradient, indicating progressive saturation of microbial N demands at high soil N. Aboveground N uptake by plants increased asymptotically with net N mineralization to a peak of approximately 35 kg N.ha(-1) x yr(-1). Aboveground net primary production per unit net N mineralization varied inversely with soil N, suggesting progressive saturation of plant N demands at high soil N. Hydrologic N losses were dominated by dissolved organic N at low-N sites, with increased nitrate loss causing a shift to dominance by nitrate at high-N sites, particularly where net nitrification exceeded plant N demands. With the exception of N mineralization patterns, our results broadly support the application of the N-saturation model developed from studies of anthropogenic N deposition to understand N cycling and saturation of plant and microbial sinks along natural soil N gradients. This convergence of behavior in unpolluted and polluted forest N cycles suggests that where future reductions in deposition to polluted sites do occur, symptoms of N saturation are most likely to persist where soil N content remains elevated.     

N fertilization effects on denitrification and N cycling in an aggrading forest.

We investigated N cycling and denitrification rates following five years of N and dolomite amendments to whole-tree harvested forest plots at the long-term soil productivity experiment in the Fernow Experimental Forest in West Virginia, USA. We hypothesized that changes in soil chemistry and nutrient cycling induced by N fertilization would increase denitrification rates and the N2O:N2 ratio. Soils from the fertilized plots had a lower pH (2.96) than control plots (3.22) and plots that received fertilizer and dolomite (3.41). There were no significant differences in soil %C or %N between treatments. Chloroform-labile microbial biomass carbon was lower in fertilized plots compared to control plots, though this trend was not significant. Extractable soil NO3- was elevated in fertilized plots on each sample date. Soil-extractable NH4+, NO3-, pH, microbial biomass carbon, and %C varied significantly by sample date suggesting important seasonal patterns in soil chemistry and N cycling. In particular, the steep decline in extractable NH4+ during the growing season is consistent with the high N demands of a regenerating forest. Net N mineralization and nitrification also varied by date but were not affected by the fertilization and dolomite treatments. In a laboratory experiment, denitrification was stimulated by NO3- additions in soils collected from all field plots, but this effect was stronger in soils from the unfertilized control plots, suggesting that chronic N fertilization has partially alleviated a NO3- limitation on denitrification rates. Dextrose stimulated denitrification only in the whole-tree-harvest soils. Denitrification enzyme activity varied by sample date and was elevated in fertilized plots for soil collected in July 2000 and June 2001. There were no detectable treatment effects on N2O or N2 flux from soils under anaerobic conditions, though there was strong temporal variation. These results suggest that whole-tree harvesting has altered the N status of these soils so they are less prone to N saturation than more mature forests. It is likely that N losses associated with the initial harvest and high N demand by aggrading vegetation is minimizing, at least temporarily, the amount of inorganic N available for nitrification and denitrification, even in the fertilized plots in this experiment.     

The ecology of energy and nutrient fluxes in hemlock forests invaded by hemlock woolly adelgid

The hemlock woolly adelgid (HWA, Adelges tsugae Annand) is currently causing a severe decline in vitality and survival of eastern hemlock in North American forests. We analyzed the effects of light HWA infestation on vertical energy and nutrient fluxes from the canopy to the forest floor. Canopy throughfall, litter lysimeters, and laboratory litter microcosms were used to examine the effects of HWA-affected and unaffected throughfall on litter type, leachate, and litter chemistry. Early in the season adelgid infestation caused higher dissolved organic carbon (DOC; +24.6%), dissolved organic nitrogen (DON; +28.5%), and K (+39.3%) fluxes and lower inorganic nitrogen fluxes (-39.8%) in throughfall and in adjacent litter solutions collected beneath infested compared to uninfested trees. Needle litter collected beneath uninfested hemlock had significantly lower N concentrations compared to needles collected beneath infested trees, while no difference in N concentrations was found in birch litter. Bacteria were significantly more abundant on hemlock and birch litter beneath infested trees, while yeasts and filamentous fungi showed no consistent response to HWA throughfall. Litter microcosms showed that less DOC was leaching from birch than from hemlock needles when exposed to HWA throughfall. Overall, NH4-N and DON leachate concentrations were higher from birch than from hemlock litter. Thus, HWA-affected throughfall leads to qualitative and quantitative differences in nitrogen export from the litter layer. The N concentration of hemlock litter did not change with time, but the N concentration in birch litter increased significantly during the course of the experiment, especially when HWA-affected throughfall was applied. We suggest a nonlinear conceptual model for the temporal and vertical transition of energy and nutrient fluxes relative to progressing HWA infestation from a pure hemlock to a birch/maple-dominated forest. Progressive needle loss and changes in needle chemistry are likely to produce a humped-shaped DOC curve, while N fluxes initially decrease as infestation continues but rise eventually with hemlock decline and immigration of hardwood species. These findings suggest that it is necessary to understand the biology and specific physiological/trophic effects of exotic pests on their hosts and associated ecosystem processes in order to decipher the temporal dynamics, direction of change, and functional consequences.     

Calcium constrains plant control over forest ecosystem nitrogen cycling

Forest ecosystem nitrogen (N) cycling is a critical controller of the ability of forests to prevent the movement of reactive N to receiving waters and the atmosphere and to sequester elevated levels of atmospheric carbon dioxide (CO2). Here we show that calcium (Ca) constrains the ability of northern hardwood forest trees to control the availability and loss of nitrogen. We evaluated soil N-cycling response to Ca additions in the presence and absence of plants and observed that when plants were present, Ca additions "tightened" the ecosystem N cycle, with decreases in inorganic N levels, potential net N mineralization rates, microbial biomass N content, and denitrification potential. In the absence of plants, Ca additions induced marked increases in nitrification (the key process controlling ecosystem N losses) and inorganic N levels. The observed "tightening" of the N cycle when Ca was added in the presence of plants suggests that the capacity of forests to absorb elevated levels of atmospheric N and CO2 is fundamentally constrained by base cations, which have been depleted in many areas of the globe by acid rain and forest harvesting.