Influences of a calcium gradient on soil inorganic nitrogen in the Adirondack Mountains, New York

Studies of the long-term impacts of acidic deposition in Europe and North America have prompted growing interest in understanding the dynamics linking the nitrogen (N) and calcium (Ca) cycles in forested watersheds. While it has been shown that increasing concentrations of nitrate (NO3-) through atmospheric deposition or through nitrification can increase Ca loss, the reciprocal effects of Ca on N transformation processes have received less attention. We studied the influence of soil Ca availability on extractable inorganic N (NO3- + NH4+) across a Ca gradient in the Adirondack Mountains, New York, USA. Our results did not show the direct Ca-N interaction that we had expected, but instead showed that exchangeable Ca coupled with soil moisture, soil organic matter, and ambient temperature accounted for 61% of the variability in extractable inorganic N across 11 sites over two growing seasons. Soil Ca concentrations were, however, positively related to sugar maple (Acer saccharum) and American basswood (Tilia americana) basal areas and negatively related to American beech (Fagus grandifolia) basal area. Based on litter chemistry differences among these tree species and reported potential N mineralization values, we suggest that the influence of Ca on soil inorganic N is through a multistep pathway: reciprocal interactions between soil Ca concentrations and species composition, which in turn affect the quality of litter available for N mineralization. If chronic soil Ca depletion continues, as reported in some forested ecosystems, potential shifts in biotic communities could result in considerable alterations of N cycling processes.     

Do evergreen and deciduous trees have different effects on net N mineralization in soil?

Evergreen and deciduous plants are widely expected to have different impacts on soil nitrogen (N) availability because of differences in leaf litter chemistry and ensuing effects on net N mineralization (N(min)). We evaluated this hypothesis by compiling published data on net N(min) rates beneath co-occurring stands of evergreen and deciduous trees. The compiled data included 35 sets of co-occurring stands in temperate and boreal forests. Evergreen and deciduous stands did not have consistently divergent effects on net N(min) rates; net N(min) beneath deciduous trees was higher when comparing natural stands (19 contrasts), but equivalent to evergreens in plantations (16 contrasts). We also compared net N(min) rates beneath pairs of co-occurring genera. Most pairs of genera did not differ consistently, i.e., tree species from one genus had higher net N(min) at some sites and lower net N(min) at other sites. Moreover, several common deciduous genera (Acer, Betula, Populus) and deciduous Quercus spp. did not typically have higher net N(min) rates than common evergreen genera (Pinus, Picea). There are several reasons why tree effects on net N(min) are poorly predicted by leaf habit and phylogeny. For example, the amount of N mineralized from decomposing leaves might be less than the amount of N mineralized from organic matter pools that are less affected by leaf litter traits, such as dead roots and soil organic matter. Also, effects of plant traits and plant groups on net N(min) probably depend on site-specific factors such as stand age and soil type.     

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。     

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.     

Leaf herbivory and decomposability in a Malaysian tropical rain forest

There is accumulating evidence that similar suites of plant traits may affect leaf palatability and leaf litter decomposability. However, the possible association between leaf herbivory and litter decomposition rates across species in species-diverse natural ecosystems such as tropical rain forests remains unexplored, despite its importance in estimating the herbivory effects on carbon and nutrient cycling of ecosystems. We found no strong association between leaf herbivory and litter decomposition rates across 40 tree species in a Malaysian tropical rain forest, even though the leaf and litter traits were tightly correlated. This is because the leaf and litter traits related to herbivory and decomposition rates in the field were inconsistent. Leaf toughness accounted for only a small part of the variation in the herbivory rate, whereas a number of litter traits (the leaf mass per area, lignin to nitrogen ratio, and condensed tannin concentration) accurately predicted the decomposition rate across species. These results suggest that herbivory rate across species may not be strongly related to single leaf traits, probably because plant-herbivore interactions in tropical rain forests are highly diverse; on the other hand, plant-decomposer interactions are less specific and can be governed by litter chemicals. We also investigated two factors, phylogeny and tree functional types, that could affect the relationship between herbivory and decomposition across species. Phylogenetic relatedness among the species did not affect the relationship between herbivory and decomposition. In contrast, when the plants were segregated according to their leaf emergence pattern, we found a significant positive relationship between herbivory and decomposition rates for continuous-leafing species. In these species, the condensed tannin to N ratios in leaves and litter were related to herbivory and decomposition rates, respectively. However, we did not observe a similar trend for synchronous-leafing species. These results suggest that the relationship between herbivory and decomposition may be more greatly affected by functional types than by phylogenetic relatedness among species. In conclusion, our results suggest that well-defended leaves are not necessarily less decomposable litter in a tropical rain forest community, implying that herbivory may not generate positive feedback for carbon and nutrient cycling in this type of ecosystem.     

Species and rotation frequency influence soil nitrogen in simplified tropical plant communities.

Among the many factors that potentially influence the rate at which nitrogen (N) becomes available to plants in terrestrial ecosystems are the identity and diversity of species composition, frequency of disturbance or stand turnover, and time. Replicated suites of investigator-designed communities afforded an opportunity to examine the effects of those factors on net N mineralization over a 12-year period. The communities consisted of large-stature perennial plants, comprising three tree species (Hyeronima alchorneoides, Cedrela odorata, and Cordia alliodora), a palm (Euterpe oleracea), and a large, perennial herb (Heliconia imbricata). Trees were grown in monoculture and in combination with the other two life-forms; tree monocultures were subjected to rotations of one or four years, or like the three-life-form systems, left uncut. The work was conducted on fertile soil in the humid lowlands of Costa Rica, a site with few abiotic constraints to plant growth. Rates of net N mineralization and nitrification were high, typically in the range of 0.2-0.8 microg x g(1) x d(-1), with net nitrification slightly higher than net mineralization, indicating preferential uptake of ammonium (NH4+) by plants and microbes. Net rates of N mineralization were about 30% lower in stands of one of the three tree species, Hyeronima, than in stands of the other two. Contrary to expectations, short-rotation management (one or four years) resulted in higher net rates of N mineralization than in uncut stands, whether the latter were composed of a single tree species or a combination of life-forms. Neither additional species richness nor replenishment of leached N augmented mineralization rates. The net rate at which N was supplied tended to be lowest in stands where demand for N was highest. Careful choice of species, coupled with low frequency of disturbance, can lead to maintenance of N within biomass and steady rates of within-system circulation, whereas pulses, whether caused by cutting and replanting or by the phenological traits of the species selected or combined, subject N supplies to leaching loss.     

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.     

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.     

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.     

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.     

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.     

Recovery of plant diversity following N cessation: effects of recruitment, litter, and elevated N cycling

Plant species richness has declined and composition shifted in response to elevated atmospheric deposition of biologically active nitrogen over much of the industrialized world. Litter thickness, litter nitrogen (N) content, and soil N mineralization rates often remain elevated long after inputs cease, clouding the prospects that plant community diversity and composition would recover should N inputs be reduced. Here we determined how N cycling, litter accumulation, and recruitment limitation influenced community recovery following cessation of long-term N inputs to prairie-like grasslands. We alleviated each of these potential inhibitors through a two-year full-factorial experiment involving organic carbon addition, litter removal, and seed addition. Seed addition had the largest effect on increasing seedling and species numbers and may be necessary to overcome long-term burial of seeds of target perennial grassland species. Litter removal increased light availability and bare sites for colonization, though it had little effect on reducing the biomass of competing neighbors or altering extractable soil N. Nonetheless, these positive influences were enough to lead to small increases in species richness within one year. We found that, although C addition quickly altered many factors assumed favorable for the target community (decreased N availability and biomass of nearby competitors, increased light and site availability), these changes were insufficient to positively impact species richness or seedling numbers over the experimental duration. However, only carbon addition had species-specific effects on the existing plant community, suggesting that its apparent limited utility may be more a result of slow recovery under ambient recruitment rather than from a lack of a restorative effect. There were dramatic interactions among treatments, with the positive effects of litter removal largely negated by carbon addition, and the positive effects of seed addition generally amplified by litter removal. It remains unclear whether each mechanism explored here will induce community recovery, but over different temporal scales. Long-term monitoring will help resolve these remaining questions. Regardless, our results suggest that reversal of species loss and compositional shifts from N deposition in prairies may be more inhibited by habitat fragmentation, recruitment limitation, and long-term suppression of fire than from continued effects of elevated N.     

Interactive effects of plant species diversity and elevated CO2 on soil biota and nutrient cycling

Terrestrial ecosystems consist of mutually dependent producer and decomposer subsystems, but not much is known on how their interactions are modified by plant diversity and elevated atmospheric CO2 concentrations. Factorially manipulating grassland plant species diversity and atmospheric CO2 concentrations for five years, we tested whether high diversity or elevated CO2 sustain larger or more active soil communities, affect soil aggregation, water dynamics, or nutrient cycling, and whether plant diversity and elevated CO2 interact. Nitrogen (N) and phosphorus (P) pools, symbiotic N2 fixation, plant litter quality, soil moisture, soil physical structure, soil nematode, collembola and acari communities, soil microbial biomass and microflora community structure (phospholipid fatty acid [PLFA] profiles), soil enzyme activities, and rates of C fluxes to soils were measured. No increases in soil C fluxes or the biomass, number, or activity of soil organisms were detected at high plant diversity; soil H2O and aggregation remained unaltered. Elevated CO2 affected the ecosystem primarily by improving plant and soil water status by reducing leaf conductance, whereas changes in C cycling appeared to be of subordinate importance. Slowed-down soil drying cycles resulted in lower soil aggregation under elevated CO2. Collembola benefited from extra soil moisture under elevated CO2, whereas other faunal groups did not respond. Diversity effects and interactions with elevated CO2 may have been absent because soil responses were mainly driven by community-level processes such as rates of organic C input and water use; these drivers were not changed by plant diversity manipulations, possibly because our species diversity gradient did not extend below five species and because functional type composition remained unaltered. Our findings demonstrate that global change can affect soil aggregation, and we advocate that soil aggregation should be considered as a dynamic property that may respond to environmental changes and feed back on other ecosystem functions.     

Tree species and mycorrhizal associations influence the magnitude of rhizosphere effects

Previous research on the effects of tree species on soil processes has focused primarily on the role of leaf litter inputs. We quantified the extent to which arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) tree species influence soil microbial activity and nutrient availability through rhizosphere effects. Rhizosphere soil, bulk soil, and fine roots were collected from 12 monospecifc plots (six AM and six ECM tree species) planted on a common soil at the Turkey Hill Plantations in Dryden, New York. Rhizosphere effects were estimated by the percentage difference between rhizosphere and bulk soil samples for several assays. Rhizosphere effects on soil microbes and their activities were significant for ECM species but in only a few cases for AM species. In AM tree species, microbial biomass, net N mineralization, and phosphatase enzyme activity in the rhizosphere were 10-12% greater than in bulk soil. In ECM tree species, rhizosphere effects for microbial biomass, C mineralization rates, net N mineralization, and phosphatase activity were 25-30% greater than bulk soil, and significantly greater than AM rhizosphere effects. The magnitude of rhizosphere effects was negatively correlated with the degree of mycorrhizal colonization in AM tree species (r = -0.83) and with fine root biomass (r = -0.88) in ECM tree species, suggesting that different factors influence rhizosphere effects in tree species forming different mycorrhizal associations. Rhizosphere effects on net N mineralization and phosphatase activity were also much greater in soils with pH < 4.3 for both AM and ECM tree species, suggesting that soil pH and its relation to nutrient availability may also influence the magnitude of rhizosphere effects. Our results support the idea that tree roots stimulate nutrient availability in the rhizosphere, and that systematic differences between AM and ECM may result in distinctive rhizosphere effects for C, N, and P cycling between AM and ECM tree species.     

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.     

Decomposers (Lumbricidae, Collembola) affect plant performance in model grasslands of different diversity

Decomposer invertebrates influence soil structure and nutrient mineralization as well as the activity and composition of the microbial community in soil and therefore likely affect plant performance and plant competition. We established model grassland communities in a greenhouse to study the interrelationship between two different functional groups of decomposer invertebrates, Lumbricidae and Collembola, and their effect on plant performance and plant nitrogen uptake in a plant diversity gradient. Common plant species of Central European Arrhenatherion grasslands were transplanted into microcosms with numbers of plant species varying from one to eight and plant functional groups varying from one to four. Separate and combined treatments with earthworms and collembolans were set up. Microcosms contained 15N labeled litter to track N fluxes into plant shoots. Presence of decomposers strongly increased total plant and plant shoot biomass. Root biomass decreased in the presence of collembolans and even more in the presence of earthworms. However, it increased when both animal groups were present. Also, presence of decomposers increased total N concentration and 15N enrichment of grasses, legumes, and small herbs. Small herbs were at a maximum in the combined treatment with earthworms and collembolans. The impact of earthworms and collembolans on plant performance strongly varied with plant functional group identity and plant species diversity and was modified when both decomposers were present. Both decomposer groups generally increased aboveground plant productivity through effects on litter decomposition and nutrient mineralization leading to an increased plant nutrient acquisition. The non-uniform effects of earthworms and collembolans suggest that functional diversity of soil decomposer animals matters and that the interactions between soil animal functional groups affect the structure of plant communities.     

Tree species control rates of free-living nitrogen fixation in a tropical rain forest

Tropical rain forests represent some of the most diverse ecosystems on earth, yet mechanistic links between tree species identity and ecosystem function in these forests remains poorly understood. Here, using free-living nitrogen (N) fixation as a model, we explore the idea that interspecies variation in canopy nutrient concentrations may drive significant local-scale variation in biogeochemical processes. Biological N fixation is the largest "natural" source of newly available N to terrestrial ecosystems, and estimates suggest the highest such inputs occur in tropical ecosystems. While patterns of and controls over N fixation in these systems remain poorly known, the data we do have suggest that chemical differences among tree species canopies could affect free-living N fixation rates. In a diverse lowland rain forest in Costa Rica, we established a series of vertical, canopy-to-soil profiles for six common canopy tree species, and we measured free-living N fixation rates and multiple aspects of chemistry of live canopy leaves, senesced canopy leaves, bulk leaf litter, and soil for eight individuals of each tree species. Free-living N fixation rates varied significantly among tree species for all four components, and independent of species identity, rates of N fixation ranged by orders of magnitude along the vertical profile. Our data suggest that variations in phosphorus (P) concentration drove a significant fraction of the observed species-specific variation in free-living N fixation rates within each layer of the vertical profile. Furthermore, our data suggest significant links between canopy and forest floor nutrient concentrations; canopy P was correlated with bulk leaf litter P below individual tree crowns. Thus, canopy chemistry may affect a suite of ecosystem processes not only within the canopy itself, but at and beneath the forest floor as well.     

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.     

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.     

Grassland establishment under varying resource availability: a test of positive and negative feedback

The traditional logic of carbon (C) and nitrogen (N) interactions in ecosystems predicts further increases or decreases in productivity (positive feedback) in response to high and low fertility in the soil, respectively; but the potential for development of feedback in ecosystems recovering from disturbance is less well understood. Furthermore, this logic has been challenged in grassland ecosystems where frequent fires or grazing may reduce the contribution of aboveground litter inputs to soil organic matter pools and nutrient supply for plant growth, relative to forest ecosystems. Further, if increases in plant productivity increase soil C content more than soil N content, negative feedback may result from increased microbial demand for N making less available for plant growth. We used a field experiment to test for feedback in an establishing grassland by comparing aboveground net primary productivity (ANPP) and belowground pools and fluxes of C and N in soil with enriched, ambient, and reduced N availability. For eight years annual N enrichment increased ANPP, root N, and root tissue quality, but root C:N ratios remained well above the threshold for net mineralization of N. There was no evidence that N enrichment increased root biomass, soil C or N accrual rates, or storage of C in total, microbial, or mineralizable pools within this time frame. However, the net nitrogen mineralization potential (NMP) rate was greater following eight years of N enrichment, and we attributed this to N saturation of the microbial biomass. Grassland developing under experimentally imposed N limitation through C addition to the soil exhibited ANPP, root biomass and quality, and net NMP rate similar to the ambient soil. Similarity in productivity and roots in the reduced and ambient N treatments was attributed to the potentially high nitrogen-use efficiency (NUE) of the dominant C4 grasses, and increasing cover of legumes over time in the C-amended soil. Thus, in a developing ecosystem, positive feedback between soil N supply and plant productivity may promote enhanced long-term N availability and override progressive N limitation as C accrues in plant and soil pools. However, experimentally imposed reduction in N availability did not feed back to reduce ANPP, possibly due to shifts in NUE and functional group composition.