High retention of 15N‐labeled nitrogen deposition in a nitrogen saturated old‐growth tropical forest

The effects of increased reactive nitrogen (N) deposition in forests depend largely on its fate in the ecosystems. However, our knowledge on the fates of deposited N in tropical forest ecosystems and its retention mechanisms is limited. Here, we report the results from the first whole ecosystem ¹⁵N labeling experiment performed in a N‐rich old‐growth tropical forest in southern China. We added ¹⁵N tracer monthly as ¹⁵NH₄¹⁵NO₃ for 1 year to control plots and to N‐fertilized plots (N‐plots, receiving additions of 50 kg N ha^?1 yr^?1 for 10 years). Tracer recoveries in major ecosystem compartments were quantified 4 months after the last addition. Tracer recoveries in soil solution were monitored monthly to quantify leaching losses. Total tracer recovery in plant and soil (N retention) in the control plots was 72% and similar to those observed in temperate forests. The retention decreased to 52% in the N‐plots. Soil was the dominant sink, retaining 37% and 28% of the labeled N input in the control and N‐plots, respectively. Leaching below 20 cm was 50 kg N ha^?1 yr^?1 in the control plots and was close to the N input (51 kg N ha^?1 yr^?1), indicating N saturation of the top soil. Nitrogen addition increased N leaching to 73 kg N ha^?1 yr^?1. However, of these only 7 and 23 kg N ha^?1 yr^?1 in the control and N‐plots, respectively, originated from the labeled N input. Our findings indicate that deposited N, like in temperate forests, is largely incorporated into plant and soil pools in the short term, although the forest is N‐saturated, but high cycling rates may later release the N for leaching and/or gaseous loss. Thus, N cycling rates rather than short‐term N retention represent the main difference between temperate forests and the studied tropical forest.     

Responses of nitrous oxide fluxes and soil nitrogen cycling to nutrient additions in montane forests along an elevation gradient in southern Ecuador

Tropical montane forests are commonly limited by N or co-limited by N and P. Projected increases in N deposition in tropical montane regions are thought to be insufficient for vegetation demand and are not therefore expected to affect soil N availability and N₂O emissions. We established a factorial N- and P-addition experiment (i.e., N, P, N + P, and control) across an elevation gradient of montane forests in Ecuador to test these hypotheses: (1) moderate rates of N and P additions are able to stimulate soil-N cycling rates and N₂O fluxes, and (2) the magnitude and timing of soil N₂O-flux responses depend on the initial nutrient status of the forest soils. Moderate rates of nutrients were added: 50 kg N ha^?1 year^?1 (in the form of urea) and 10 kg P ha^?1 year^?1 (in the form of NaH₂PO ₄ ^. 2H₂O) split in two equal applications. We tested the hypotheses by measuring changes in net rates of soil–N cycling and N₂O fluxes during the first 2 years (2008–2009) of nutrient manipulation in an old-growth premontane forest at 1,000 m, growing on a Cambisol soil with no organic layer, in an old-growth lower montane forest at 2,000 m, growing on a Cambisol soil with an organic layer, and an old-growth upper montane rainforest at 3,000 m, growing on a Histosol soil with a thick organic layer. Among the control plots, net nitrification rates were largest at the 1,000-m site whereas net nitrification was not detectable at the 2,000- and 3,000-m sites. The already large net nitrification at the 1,000-m site was not affected by nutrient additions, but net nitrification became detectable at the 2,000- and 3000-m sites after the second year of N and N + P additions. N₂O emissions increased rapidly following N and N + P additions at the 1,000-m site whereas only smaller increases occurred at the 2,000- and 3,000-m sites during the second year of N and N + P additions. Addition of P alone had no effect on net rates of soil N cycling and N₂O fluxes at any elevation. Our results showed that the initial soil N status, which may also be influenced by presence or absence of organic layer, soil moisture and temperature as encompassed by the elevation gradient, is a good indicator of how soil N cycling and N₂O fluxes may respond to future increases in nutrient additions.     

Nitrogen Addition Shapes Soil Phosphorus Availability in Two Reforested Tropical Forests in Southern China

Scant information is available on how soil phosphorus (P) availability responds to atmospheric nitrogen (N) deposition, especially in the tropical zones. This study examined the effect of N addition on soil P availability, and compared this effect between forest sites of contrasting land‐use history. Effects of N addition on soil properties, litterfall production, P release from decomposing litter, and soil P availability were studied in a disturbed (reforested pine forest with previous understory vegetation and litter harvesting) and a rehabilitated (reforested mixed pine/broadleaf forest with no understory vegetation and litter harvesting) tropical forest in southern China. Experimental N‐treatments (above ambient) were the following: Control (no N addition), N50 (50 kg N ha^?1 yr^?1), and N100 (100 kg N ha^?1 yr^?1). Results indicated that N addition significantly decreased soil P availability in the disturbed forest. In the rehabilitated forest, however, soil P availability was significantly increased by N addition. Decreases in soil P availability may be correlated with decreases in rates of P release from decomposing litter in the N‐treated plots, whereas the increase in soil P availability was correlated with an increase in litterfall production. Our results suggest that response of soil P availability to N deposition in the reforested tropical forests in southern China may vary greatly with temporal changes in tree species composition and soil nutrient status, caused by different land‐use practices.     

Major changes in forest carbon and nitrogen cycling caused by declining sulphur deposition

Sulphur (S) and nitrogen (N) deposition are important drivers of the terrestrial carbon (C) and N cycling. We analyzed changes in C and N pools in soil and tree biomass at a highly acidified spruce site in the Czech Republic during a 15 year period. Total S deposition decreased from 5 to 1.1 g m^?2 yr^?1 between 1995 and 2009, whereas bulk N deposition did not change. Over the same period, C and N pools in the Oa horizon declined by 116 g C and 4.2 g N m^?2 yr^?1, a total decrease of 47% and 42%, respectively. This loss of C and N probably originated from organic matter (OM) that had accumulated during the period of high acid deposition when litter decomposition was suppressed. The loss of OM from the Oa horizon coincided with a substantial leaching (1.3 g N m^?2 yr^?1 at 90 cm) in the 1990s to almost no leaching (<0.02 g N m^?2 yr^?1) since 2006. Forest floor net N mineralization also decreased. This had consequences for spruce needle N concentration (from 17.1 to 11.4 mg kg^?1 in current needles), an increase in litterfall C/N ratio (from 51 to 63), and a significant increase in the Oi + Oe horizon C/N ratio (from 23.4 to 27.3) between 1994 and 2009/2010. Higher forest growth and lower canopy defoliation was observed in the 2000s compared to the 1990s. Our results demonstrate that reducing S deposition has had a profound impact on forest organic matter cycling, leading to a reversal of historic ecosystem N enrichment, cessation of nitrate leaching, and a major loss of accumulated organic soil C and N stocks. These results have major implications for our understanding of the controls on both N saturation and C sequestration in forests, and other ecosystems, subjected to current or historic S deposition.     

The influence of tree species on canopy soil nutrient status in a tropical lowland wet forest in Costa Rica

The canopy is host to a large percentage of the flora and fauna in tropical wet forests and is distinct from the forest floor in plant richness, soil type and microclimate. In this study, we examined the influence of tree species and season on soil nutrient cycling processes in canopy soils of four tree species common to Costa Rican wet forests. We also compared the canopy soils to the associated forest floor mineral soils. Both tree species and season had strong effects on canopy soil nutrients and processes. Canopy soils from trees with high litter lignin concentrations had higher net N-mineralization rates and higher dissolved inorganic N concentrations than those with low lignin concentrations. During the dry season, net N-immobilization occurred and dissolved organic and inorganic N and available P concentrations were significantly higher than during the wet season. Overall, canopy soils had higher N levels and higher fungi + bacteria richness than forest floor mineral soils. The differences in canopy soil properties observed among tree species indicates that these species have distinct N cycles that reflect differences in both soil origin and biological controls.     

Topography strongly affects atmospheric deposition and canopy exchange processes in different types of wet lowland rainforest,Southwest Costa Rica

Bulk precipitation and throughfall were collected in a wet lowland rainforest in SW Costa Rica on an event basis to allow modelling the contributions of dry deposition and canopy exchange to nutrient inputs and internal cycling of nutrients. Estimates based on bulk precipitation underestimated total atmospheric deposition to tropical rainforests by up to 10-fold ignoring the contributions of dry deposition. Canopy exchange contributed most of the aboveground inputs to the forest soil of Na⁺, about half for K⁺, 10% for P and Mg²⁺ and negligible for N, C and other elements. Tree species composition did not account for the differences found in net throughfall between forest sites, and vegetation structure (plant area index) had only a small effect on net throughfall. Forest regrowth affected net throughfall through reduced soil fertility and differences in leaf traits. Topography most significantly affected net throughfall via increased dry deposition at sites of higher elevation and via soil fertility and increased canopy exchange at down slope sites.     

Canopy soil greenhouse gas dynamics in response to indirect fertilization across an elevation gradient of tropical montane forests

Canopy soils can significantly contribute to aboveground labile biomass, especially in tropical montane forests. Whether they also contribute to the exchange of greenhouse gases is unknown. To examine the importance of canopy soils to tropical forest‐soil greenhouse gas exchange, we quantified gas fluxes from canopy soil cores along an elevation gradient with 4 yr of nutrient addition to the forest floor. Canopy soil contributed 5–12 percent of combined (canopy + forest floor) soil CO₂ emissions but CH₄ and N₂O fluxes were low. At 2000 m, phosphorus decreased CO₂ emissions (>40%) and nitrogen slightly increased CH₄ uptake and N₂O emissions. Our results show that canopy soils may contribute significantly to combined soil greenhouse gas fluxes in montane regions with high accumulations of canopy soil. We also show that changes in fluxes could occur with chronic nutrient deposition.     

On the tracks of Nitrogen deposition effects on temperate forests at their southern European range – an observational study from Italy

We studied forest monitoring data collected at permanent plots in Italy over the period 2000–2009 to identify the possible impact of nitrogen (N) deposition on soil chemistry, tree nutrition and growth. Average N throughfall (N‐NO₃+N‐NH₄) ranged between 4 and 29 kg ha^?1 yr^?1, with Critical Loads (CLs) for nutrient N exceeded at several sites. Evidence is consistent in pointing out effects of N deposition on soil and tree nutrition: topsoil exchangeable base cations (BCE) and pH decreased with increasing N deposition, and foliar nutrient N ratios (especially N : P and N : K) increased. Comparison between bulk openfield and throughfall data suggested possible canopy uptake of N_, levelling out for bulk deposition >4–6 kg ha^?1 yr^?1. Partial Least Square (PLS) regression revealed that ‐ although stand and meteorological variables explained the largest portion of variance in relative basal area increment (BAI_rel 2000–2009) ‐ N‐related predictors (topsoil BCE, C : N, pH; foliar N‐ratios; N deposition) nearly always improved the BAI_rel model in terms of variance explained (from 78.2 to 93.5%) and error (from 2.98 to 1.50%). N deposition was the strongest predictor even when stand, management and atmosphere‐related variables (meteorology and tropospheric ozone) were accounted for. The maximal annual response of BAI_rel was estimated at 0.074–0.085% for every additional kgN. This corresponds to an annual maximal relative increase of 0.13–0.14% of carbon sequestered in the above‐ground woody biomass for every additional kgN, i.e. a median value of 159 kgC per kgN ha^?1 yr^?1 (range: 50–504 kgC per kgN, depending on the site). Positive growth response occurred also at sites where signals of possible, perhaps recent N saturation were detected. This may suggest a time lag for detrimental N effects, but also that, under continuous high N input, the reported positive growth response may be not sustainable in the long‐term.     

Soil moisture effects on gross nitrification differ between adjacent grassland and forested soils in central Alberta, Canada

Background and aims

Changes in soil moisture availability seasonally and as a result of climatic variability would influence soil nitrogen (N) cycling in different land use systems. This study aimed to understand mechanisms of soil moisture availability on gross N transformation rates.

Methods

A laboratory incubation experiment was conducted to evaluate the effects of soil moisture content (65 vs. 100% water holding capacity, WHC) on gross N transformation rates using the ¹⁵N tracing technique (calculated by the numerical model FLUAZ) in adjacent grassland and forest soils in central Alberta, Canada.

Results

Gross N mineralization and gross NH ₄ ⁺ immobilization rates were not influenced by soil moisture content for both soils. Gross nitrification rates were greater at 100 than at 65% WHC only in the forest soil. Denitrification rates during the 9 days of incubation were 2.47 and 4.91 mg N kg^-1 soil d^-1 in the grassland and forest soils, respectively, at 100% WHC, but were not different from zero at 65% WHC. In the forest soil, both the ratio of gross nitrification to gross NH ₄ ⁺ immobilization rates (N/IA) and cumulative N₂O emission were lower in the 65 than in the 100% WHC treatment, while in the grassland soil, the N/IA ratio was similar between the two soil moisture content treatments but cumulative N₂O emission was lower at 65% WHC.

Conclusions

The effect of soil moisture content on gross nitrification rates differ between forest and grassland soils and decreasing soil moisture content from 100 to 65% WHC reduced N₂O emissions in both soils.     

Nitrogen deposition contributes to soil acidification in tropical ecosystems

Elevated anthropogenic nitrogen (N) deposition has greatly altered terrestrial ecosystem functioning, threatening ecosystem health via acidification and eutrophication in temperate and boreal forests across the northern hemisphere. However, response of forest soil acidification to N deposition has been less studied in humid tropics compared to other forest types. This study was designed to explore impacts of long‐term N deposition on soil acidification processes in tropical forests. We have established a long‐term N‐deposition experiment in an N‐rich lowland tropical forest of Southern China since 2002 with N addition as NH₄NO₃ of 0, 50, 100 and 150 kg N ha^?1 yr^?1. We measured soil acidification status and element leaching in soil drainage solution after 6‐year N addition. Results showed that our study site has been experiencing serious soil acidification and was quite acid‐sensitive showing high acidification (pH_(H2O)<4.0), negative water‐extracted acid neutralizing capacity (ANC) and low base saturation (BS,< 8%) throughout soil profiles. Long‐term N addition significantly accelerated soil acidification, leading to depleted base cations and decreased BS, and further lowered ANC. However, N addition did not alter exchangeable Al³⁺, but increased cation exchange capacity (CEC). Nitrogen addition‐induced increase in SOC is suggested to contribute to both higher CEC and lower pH. We further found that increased N addition greatly decreased soil solution pH at 20 cm depth, but not at 40 cm. Furthermore, there was no evidence that Al³⁺ was leaching out from the deeper soils. These unique responses in tropical climate likely resulted from: exchangeable H⁺ dominating changes of soil cation pool, an exhausted base cation pool, N‐addition stimulating SOC production, and N saturation. Our results suggest that long‐term N addition can contribute measurably to soil acidification, and that shortage of Ca and Mg should receive more attention than soil exchangeable Al in tropical forests with elevated N deposition in the future.     

Substrate, climate, and land use controls over soil N dynamics and N-oxide emissions in Borneo

Nitrogen (N) enrichment of tropical ecosystems is likely to increase with rapid industrial and agricultural development, but the ecological consequences of N additions in these systems are not well understood. We measured soil N- oxide emissions and N transformations in primary rain forest ecosystems at four elevations and across two substrate types on Mt. Kinabalu, Borneo, before and after short-term experimental N additions. We also measured N pools and fluxes across a land use gradient of primary forest, burned secondary forest, and fertilized agriculture. Background soil N₂O and NO emissions in primary forest decreased with elevation, and soils derived from sedimentary substrates had larger pools of inorganic N, rates of nitrification, and N-oxide fluxes than ultrabasic soils when there were significant differences between substrate types. N-oxide emissions after N additions and background rates of nitrification were low in all soils derived from ultrabasic substrates compared to sedimentary substrates, even at lowland sites supporting, diverse Dipterocarp forests growing on morphologically similar Oxisols. Rates of potential nitrification were good predictors of N-oxide emissions after N additions. N₂O and NO fluxes were largest at low elevations and on sedimentary-derived soils compared to ultrabasic-derived soils, even at the smallest addition of N, 15kgNha^–1. Because current methods of soil classification do not explicitly characterize a number of soil chemical properties important to nutrient cycling, the use of soil maps to extrapolate biogeochemical processes to the region or globe may be limited in its accuracy and usefulness. In agricultural systems, management practices were more important than substrate type in controlling N-oxide emissions and soil N cycling. N-oxide fluxes from agricultural fields were more than an order of magnitude greater than from primary forests on the same substrate type and at the same elevation. As primary forests are cleared for intensive agriculture, soil N₂O and NO emissions are likely to far exceed those from the most N-saturated tropical forest ecosystems. This study highlights the inter-dependence of climate, substrate age, N deposition, and land-use practices determining N cycling and N-oxide emissions in humid tropical regions.     

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.     

Litter Decomposition Within the Canopy and Forest Floor of Three Tree Species in a Tropical Lowland Rain forest,Costa Rica

The rain forest canopy hosts a large percentage of the world's plant biodiversity, which is maintained, in large part, by internal nutrient cycling. This is the first study to examine the effects of site (canopy, forest floor) and tree species (Dipteryx panamensis, Lecythis ampla, Hyeronima alchorneoides) on decay rates of a common substrate and in situ leaf litter in a tropical forest in Costa Rica. Decay rates were slower for both substrates within the canopy than on the forest floor. The slower rate of mass loss of the common substrate in the canopy was due to differences in microclimate between sites. Canopy litter decay rates were negatively correlated with litter lignin:P ratios, while forest floor decay rates were negatively correlated with lignin concentrations, indicating that the control of litter decay rates in the canopy is P availability while that of the forest floor is carbon quality. The slower cycling rates within the canopy are consistent with lower foliar nutrient concentrations of epiphytes compared with forest floor-rooted plants. Litter decay rates, but not common substrate decay rates, varied among tree species. The lack of variation in common substrate decay among tree species eliminated microclimatic variation as a possible cause for differences in litter decay and points to variation in litter quality, nutrient availability and decomposer community of tree species as the causal factors. The host tree contribution to canopy nutrient cycling via litter quality and inputs may influence the quality and quantity of canopy soil resources.     

Changes in nitrogen cycling and retention processes in soils under spruce forests along a nitrogen enrichment gradient in Germany

A network of long-term monitoring sites on nitrogen (N) input and output of forests across Germany showed that a number of Germany's forests are subject to or are experiencing N saturation and that spruce (Picea abies) stands have high risk. Our study was aimed at (1) quantifying the changes in gross rates of microbial N cycling and retention processes in forest soils along an N enrichment gradient and (2) relating the changes in soil N dynamics to N losses. We selected spruce sites representing an N enrichment gradient (indicated by leaching : throughfall N ratios) ranging from 0.04–0.13 (low N),≤0.26 (intermediate N enrichment) to≥0.42 (highly N enriched). To our knowledge, our study is the first to report on mechanistic changes in gross rates of soil N cycling and abiotic NO₃⁻ retention under ambient N enrichment gradient. Gross N mineralization, NH₄⁺ immobilization, gross nitrification, and NO₃⁻ immobilization rates increased up to intermediate N enrichment level and somewhat decreased at highly N-enriched condition. The turnover rates of NH₄⁺ and microbial N pools increased while the turnover rates of the NO₃⁻ pool decreased across the N enrichment gradient. Abiotic immobilization of NH₄⁺ did not differ across sites and was lower than that of NO₃⁻. Abiotic NO₃⁻ immobilization decreased across the N enrichment gradient. Microbial assimilation and turnover appeared to contribute largely to the retention of NH₄⁺. The increasing NO₃⁻ deposition and decreasing turnover rates of the NO₃⁻ pool, combined with decreasing abiotic NO₃⁻ retention, possibly contributed to increasing NO₃⁻ leaching and gaseous emissions across the N enrichment gradient. The empirical relationships of changes in microbial N cycling across the N enrichment gradient may be integrated in models used to predict responses of forest ecosystems (e.g. spruce) to increasing N deposition.     

Biological Nitrogen Fixation in Two Tropical Forests: Ecosystem-Level Patterns and Effects of Nitrogen Fertilization

Humid tropical forests are often characterized by large nitrogen (N) pools, and are known to have large potential N losses. Although rarely measured, tropical forests likely maintain considerable biological N fixation (BNF) to balance N losses. We estimated inputs of N via BNF by free-living microbes for two tropical forests in Puerto Rico, and assessed the response to increased N availability using an on-going N fertilization experiment. Nitrogenase activity was measured across forest strata, including the soil, forest floor, mosses, canopy epiphylls, and lichens using acetylene (C₂H₂) reduction assays. BNF varied significantly among ecosystem compartments in both forests. Mosses had the highest rates of nitrogenase activity per gram of sample, with 11 ± 6 nmol C₂H₂ reduced/g dry weight/h (mean ± SE) in a lower elevation forest, and 6 ± 1 nmol C₂H₂/g/h in an upper elevation forest. We calculated potential N fluxes via BNF to each forest compartment using surveys of standing stocks. Soils and mosses provided the largest potential inputs of N via BNF to these ecosystems. Summing all components, total background BNF inputs were 120 ± 29 μg N/m²/h in the lower elevation forest, and 95 ± 15 μg N/m²/h in the upper elevation forest, with added N significantly suppressing BNF in soils and forest floor. Moisture content was significantly positively correlated with BNF rates for soils and the forest floor. We conclude that BNF is an active biological process across forest strata for these tropical forests, and is likely to be sensitive to increases in N deposition in tropical regions.     

Net nitrogen mineralization and net nitrification along a tropical forest-to-pasture chronosequence

Soil inorganic nitrogen pools, net mineralization and net nitrification rates were compared during the dry season along a chronosequence of upland (terra firme) forest, 3-, 9- and 20-year-old pastures in the western Brazilian Amazon Basin state of Rondônia to investigate the influence of forest conversion to pasture on soil nitrogen cycles. Surface soil (0 to 10 cm) from forest had larger extractable inorganic nitrogen pools than pasture soils. In the forest, NO ₃ ^– pools equaled or exceeded NH ₄ ⁺ pools, while pasture inorganic N pools consisted almost exclusively of NH ₄ ⁺ . Rates of net N mineralization and net nitrification in seven -day laboratory incubations were higher in the seven - day forest than in the pastures. Net N mineralization rates did not differ significantly among different-aged pastures, but net nitrification rates were significantly lower in the 20-year-old pasture. Higher net N mineralization and net nitrification rates were measured in laboratory and in situ incubations of sieved soil, compared with in situ incubations of intact soil cores. Rates calculated in seven-day incubations were higher than determined by longer incubations. Sieving may increase N mineralization and/or decrease N immobilization compared with intact cores. We concluded that 7-day laboratory incubation of sieved soil was the most useful index for comparing N availability across the chronosequence of forest and pasture sites. High net nitrification rates in forest soils suggest a potential for NO ₃ ^– losses either through leaching or gaseous emissions.     

Contrasting effects of low and high nitrogen additions on soil CO2 flux components and ectomycorrhizal fungal sporocarp production in a boreal forest

Nitrogen (N) added through atmospheric deposition or as fertilizer to boreal and temperate forests reduces both soil decomposer activity (heterotrophic respiration) and the activity of roots and mycorrhizal fungi (autotrophic respiration). However, these negative effects have been found in studies that applied relatively high levels of N, whereas the responses to ambient atmospheric N deposition rates are still not clear. Here, we compared an unfertilized control boreal forest with a fertilized forest (100 kg N ha^?1 yr^?1) and a forest subject to N‐deposition rates comparable to those in Central Europe (20 kg N ha^?1 yr^?1) to investigate the effects of N addition rate on different components of forest floor respiration and the production of ectomycorrhizal fungal sporocarps. Soil collars were used to partition heterotrophic (R_h) and autotrophic (R_a) respiration, which was further separated into respiration by tree roots (R_tr) and mycorrhizal hyphae (R_m). Total forest floor respiration was twice as high in the low N plot compared to the control, whereas there were no differences between the control and high N plot. There were no differences in R_h respiration among plots. The enhanced forest floor respiration in the low N plot was, therefore, the result of increased R_a respiration, with an increase in R_tr respiration, and a doubling of R_m respiration. The latter was corroborated by a slightly greater ectomycorrhizal (EM) fungal sporocarp production in the low N plot as compared to the control plot. In contrast, EM fungal sporocarp production was nearly eliminated, and R_m respiration severely reduced, in the high N plot, which resulted in significantly lower R_a respiration. We thus found a nonlinear response of the R_a components to N addition rate, which calls for further studies of the quantitative relations among N addition rate, plant photosynthesis and carbon allocation, and the function of EM fungi.     

Imaging spectroscopy reveals the effects of topography and logging on the leaf chemistry of tropical forest canopy trees

Logging, pervasive across the lowland tropics, affects millions of hectares of forest, yet its influence on nutrient cycling remains poorly understood. One hypothesis is that logging influences phosphorus (P) cycling, because this scarce nutrient is removed in extracted timber and eroded soil, leading to shifts in ecosystem functioning and community composition. However, testing this is challenging because P varies within landscapes as a function of geology, topography and climate. Superimposed upon these trends are compositional changes in logged forests, with species with more acquisitive traits, characterized by higher foliar P concentrations, more dominant. It is difficult to resolve these patterns using traditional field approaches alone. Here, we use airborne light detection and ranging‐guided hyperspectral imagery to map foliar nutrient (i.e. P, nitrogen [N]) concentrations, calibrated using field measured traits, over 400 km² of northeastern Borneo, including a landscape‐level disturbance gradient spanning old‐growth to repeatedly logged forests. The maps reveal that canopy foliar P and N concentrations decrease with elevation. These relationships were not identified using traditional field measurements of leaf and soil nutrients. After controlling for topography, canopy foliar nutrient concentrations were lower in logged forest than in old‐growth areas, reflecting decreased nutrient availability. However, foliar nutrient concentrations and specific leaf area were greatest in relatively short patches in logged areas, reflecting a shift in composition to pioneer species with acquisitive traits. N:P ratio increased in logged forest, suggesting reduced soil P availability through disturbance. Through the first landscape scale assessment of how functional leaf traits change in response to logging, we find that differences from old‐growth forest become more pronounced as logged forests increase in stature over time, suggesting exacerbated phosphorus limitation as forests recover.     

Large Canopy Exchange Fluxes of Inorganic and Organic Nitrogen and Preferential Retention of Nitrogen by Epiphytes in a Tropical Lowland Rainforest

Little is known about how tropical forest canopies interact with atmospheric nitrogen deposition and how this affects the internal nutrient dynamics and the processing of external nutrient inputs. The objectives of this study therefore were (1) to investigate gross and net canopy nitrogen (N) fluxes (retention and leaching) and (2) the effect of canopy components on net canopy N retention. Tracers were applied on detached branches in a tropical wet lowland rainforest, Costa Rica. A novel ¹⁵N pool dilution method showed that gross canopy fluxes (retention and leaching) of NO₃ ^?, NH₄ ⁺, and dissolved organic nitrogen (DON) were remarkably higher than net throughfall fluxes. Gross fluxes of NH₄ ⁺ and NO₃ ^? resulted in a negligible net flux whereas DON showed net uptake by the canopy. The highest quantity of ¹⁵N was recovered in epiphytic bryophytes (16.4%) although the largest biomass fraction was made up of leaves. The study demonstrates that tracer applications allow investigation of the dynamic and complex canopy exchange processes and that epiphytic communities play a major role in solute fluxes in tree canopies and therefore in the nutrient dynamics of tropical rain forests.     

Nitrogen cycling in forest soils across climate gradients in Eastern China

A ¹⁵N tracing study was carried out to investigate the potential gross nitrogen (N) dynamics in thirteen forest soils in Eastern China ranging from temperate to tropical zones (five coniferous forests, six deciduous broad-leaf forests, one temperate mixed forest, one evergreen broad-leaf forests ecosystems), and to identify the major controlling factors on N cycling in these forest ecosystems. The soil pH ranged from 4.3 to 7.9 and soil organic carbon (SOC) ranged from 6.6 g?kg^?1 to 83.0 g?kg^?1. The potential gross N transformation rates were quantified by ¹⁵N tracing studies where either the ammonium or nitrate pools were ¹⁵N labeled in parallel treatments. Gross mineralization rates ranged from 0.915 μg N g^?1 soil day^?1 to 2.718 μg N g^?1 soil day^?1 in the studied forest soils. The average contribution of labile organic-N (M _Nlab ) to total gross mineralization (M _Nrec +M _Nlab ) was 86% (58% to 99%), indicating that turnover of labile organic N plays a dominant role in the studied forest ecosystems. The gross mineralization rates in coniferous forest soils were significantly lower (ranging between 0.915 and 1.228 μg N g^?1 soil day^?1) compared to broad-leaf forest soils (ranging from 1.621 to 2.718 μg N g^?1 soil day^?1) (p?<?0.01). Thus, the dominant vegetation may play an important role in regulating soil N mineralization. Nitrate production (nitrification) occurred via two pathways, oxidation of NH ₄ ⁺ and organic N the forest soils. Correlations with soil pH indicated that this is a key factor controlling the oxidation of NH ₄ ⁺ and organic N in theses forest ecosystems. NH ₄ ⁺ oxidation decreased with a decline in pH while organic N oxidation increased. The climatic conditions (e.g. moisture status) at the various sites governed the NO ₃ ^? -N consumption processes (dissimilatory NO ₃ ^? reduction to NH ₄ ⁺ (DNRA) or immobilization of NO ₃ ^? ). Total NO ₃ ^? consumption and the proportion of total NO ₃ ^? consumption to total NO ₃ ^? production decreased with an increase in the drought index of ecosystems, showing that strong interactions appear to exist between climatic condition (e.g. the drought index), N mineralization and the rate of DNRA. Interactions between vegetation, climatic conditions govern internal N cycling in these forests soils.