Effects of nitrogen additions on above- and belowground carbon dynamics in two tropical forests

Anthropogenic nitrogen (N) deposition is increasing rapidly in tropical regions, adding N to ecosystems that often have high background N availability. Tropical forests play an important role in the global carbon (C) cycle, yet the effects of N deposition on C cycling in these ecosystems are poorly understood. We used a field N-fertilization experiment in lower and upper elevation tropical rain forests in Puerto Rico to explore the responses of above- and belowground C pools to N addition. As expected, tree stem growth and litterfall productivity did not respond to N fertilization in either of these N-rich forests, indicating a lack of N limitation to net primary productivity (NPP). In contrast, soil C concentrations increased significantly with N fertilization in both forests, leading to larger C stocks in fertilized plots. However, different soil C pools responded to N fertilization differently. Labile (low density) soil C fractions and live fine roots declined with fertilization, while mineral-associated soil C increased in both forests. Decreased soil CO₂ fluxes in fertilized plots were correlated with smaller labile soil C pools in the lower elevation forest (R² = 0.65, p < 0.05), and with lower live fine root biomass in the upper elevation forest (R² = 0.90, p < 0.05). Our results indicate that soil C storage is sensitive to N deposition in tropical forests, even where plant productivity is not N-limited. The mineral-associated soil C pool has the potential to respond relatively quickly to N additions, and can drive increases in bulk soil C stocks in tropical forests.     

Microbial activity and soil respiration under nitrogen addition in Alaskan boreal forest

Climate warming could increase rates of soil organic matter turnover and nutrient mineralization, particularly in northern high‐latitude ecosystems. However, the effects of increasing nutrient availability on microbial processes in these ecosystems are poorly understood. To determine how soil microbes respond to nutrient enrichment, we measured microbial biomass, extracellular enzyme activities, soil respiration, and the community composition of active fungi in nitrogen (N) fertilized soils of a boreal forest in central Alaska. We predicted that N addition would suppress fungal activity relative to bacteria, but stimulate carbon (C)‐degrading enzyme activities and soil respiration. Instead, we found no evidence for a suppression of fungal activity, although fungal sporocarp production declined significantly, and the relative abundance of two fungal taxa changed dramatically with N fertilization. Microbial biomass as measured by chloroform fumigation did not respond to fertilization, nor did the ratio of fungi : bacteria as measured by quantitative polymerase chain reaction. However, microbial biomass C : N ratios narrowed significantly from 16.0 ± 1.4 to 5.2 ± 0.3 with fertilization. N fertilization significantly increased the activity of a cellulose‐degrading enzyme and suppressed the activities of protein‐ and chitin‐degrading enzymes but had no effect on soil respiration rates or ¹⁴C signatures. These results indicate that N fertilization alters microbial community composition and allocation to extracellular enzyme production without affecting soil respiration. Thus, our results do not provide evidence for strong microbial feedbacks to the boreal C cycle under climate warming or N addition. However, organic N cycling may decline due to a reduction in the activity of enzymes that target nitrogenous compounds.     

Variable Responses of Lowland Tropical Forest Nutrient Status to Fertilization and Litter Manipulation

Predicting future impacts of anthropogenic change on tropical forests requires a clear understanding of nutrient constraints on productivity. We compared experimental fertilization and litter manipulation treatments in an old-growth lowland tropical forest to distinguish between the effects of inorganic nutrient amendments and changes in nutrient cycling via litterfall. We measured the changes in soil and litter nutrient pools, litterfall, and fine root biomass in plots fertilized with nitrogen (N), phosphorus (P), or potassium (K), and in litter addition and litter removal treatments during 7 years. Soil inorganic N and litter N increased in double-litter plots but not in N-fertilized plots. Conversely, litter P and soil pools of P and K increased in fertilized plots but not in the double-litter plots. Soil and litter pools of N and K decreased in the no-litter plots. Changes in litterfall with added nutrients or litter were only marginally significant, but fine root biomass decreased with both the litter and the K addition. Differences between the two experiments are mostly attributable to the coupled cycling of carbon and nutrients in litter. Increased nutrient inputs in litter may improve plant uptake of some nutrients compared to fertilization with similar amounts. The litter layer also appears to play a key role in nutrient retention. We discuss our findings in the context of possible impacts of anthropogenic change on tropical forests.     

Effects of nutrient additions on ecosystem carbon cycle in a Puerto Rican tropical wet forest

Wet tropical forests play a critical role in global ecosystem carbon (C) cycle, but C allocation and the response of different C pools to nutrient addition in these forests remain poorly understood. We measured soil organic carbon (SOC), litterfall, root biomass, microbial biomass and soil physical and chemical properties in a wet tropical forest from May 1996 to July 1997 following a 7‐year continuous fertilization. We found that although there was no significant difference in total SOC in the top 0–10 cm of the soils between the fertilization plots (5.42±0.18 kg m^?2) and the control plots (5.27±0.22 kg m^?2), the proportion of the heavy‐fraction organic C in the total SOC was significantly higher in the fertilized plots (59%) than in the control plots (46%) (P<0.05). The annual decomposition rate of fertilized leaf litter was 13% higher than that of the control leaf litter. We also found that fertilization significantly increased microbial biomass (fungi+bacteria) with 952±48 mg kg^?1soil in the fertilized plots and 755±37 mg kg^?1soil in the control plots. Our results suggest that fertilization in tropical forests may enhance long‐term C sequestration in the soils of tropical wet forests.     

Nitrogen cycling in canopy soils of tropical montane forests responds rapidly to indirect N and P fertilization

Although the canopy can play an important role in forest nutrient cycles, canopy‐based processes are often overlooked in studies on nutrient deposition. In areas of nitrogen (N) and phosphorus (P) deposition, canopy soils may retain a significant proportion of atmospheric inputs, and also receive indirect enrichment through root uptake followed by throughfall or recycling of plant litter in the canopy. We measured net and gross rates of N cycling in canopy soils of tropical montane forests along an elevation gradient and assessed indirect effects of elevated nutrient inputs to the forest floor. Net N cycling rates were measured using the buried bag method. Gross N cycling rates were measured using ¹⁵N pool dilution techniques. Measurements took place in the field, in the wet and dry season, using intact cores of canopy soil from three elevations (1000, 2000 and 3000 m). The forest floor had been fertilized biannually with moderate amounts of N and P for 4 years; treatments included control, N, P, and N + P. In control plots, gross rates of NH₄⁺ transformations decreased with increasing elevation; gross rates of NO₃^? transformations did not exhibit a clear elevation trend, but were significantly affected by season. Nutrient‐addition effects were different at each elevation, but combined N + P generally increased N cycling rates at all elevations. Results showed that canopy soils could be a significant N source for epiphytes as well as contributing up to 23% of total (canopy + forest floor) mineral N production in our forests. In contrast to theories that canopy soils are decoupled from nutrient cycling in forest floor soil, N cycling in our canopy soils was sensitive to slight changes in forest floor nutrient availability. Long‐term atmospheric N and P deposition may lead to increased N cycling, but also increased mineral N losses from the canopy soil system.     

Anthropogenic N Deposition Increases Soil C Storage by Decreasing the Extent of Litter Decay: Analysis of Field Observations with an Ecosystem Model

Recent meta-analyses of experimental studies simulating increased anthropogenic nitrogen (N) deposition in forests reveal greater soil carbon (C) storage under elevated levels of atmospheric N deposition. However, these effects have not yet been included in ecosystem-scale models of soil C and N cycling and it is unclear whether increased soil C storage results from slower decomposition rates or a reduced extent of decomposition (for example, an increase in the amount of litter entering slowly decaying humus pools). To test these alternatives, we conducted a meta-analysis of litter decomposition data. We then used the results from our meta-analysis to model C and N cycling in four sugar maple forests in Michigan using an ecosystem process model (TRACE). We compared model results testing our alternative hypotheses to field data on soil C storage from a 17-year N deposition experiment. Using data from published litter decomposition studies in forests, we determined that, on average, exogenous N inputs decreased lignin decomposition rates by 30% and increased cellulose decomposition by 9%. In the same set of litter decomposition studies increased exogenous N availability increased the amount of litter entering slowly decaying humus pools in a manner significantly related to the lignocellulose index of decaying litter. Incorporating changes to decomposition rates in TRACE did not accurately reproduce greater soil C storage observed in our field study with experimentally elevated N deposition. However, when changes in the extent of decomposition were incorporated in TRACE, the model produced increased soil C storage by increasing the amount of litter entering the humus pool and accurately represented C storage in plant and soil pools under experimental N deposition. Our modeling results and meta-analysis indicate that the extent of litter decay as humus is formed, rather than slower rates of litter decay, is likely responsible for the accumulation of organic matter, and hence soil C storage, under experimental N deposition. This effect should be incorporated in regional to global-scale models simulating the C balance of forest ecosystems in regions receiving elevated N deposition.     

Nutrient availability controls the impact of mammalian herbivores on soil carbon and nitrogen pools in grasslands

Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature – herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local‐scale herbivory, and its interaction with nutrient enrichment and climate, within global‐scale models to better predict land–atmosphere interactions under future climate change.     

Responses of soil microbial biomass and enzymatic activities to different forms of organic nitrogen deposition in the subtropical forests in East China

Numerous studies reported that inorganic nitrogen (N) deposition strongly affected forest ecosystems. However, organic N is also an important component of atmospheric N deposition. The influence of organic N deposition on soil microbial biomass and extracellular enzymatic activities (EEA) in subtropical forests remains unclear. Coniferous forest (CF) and broad-leaved forest (BF) were chosen from the Zijin Mountain in China. Five forms of organic N (urea, glycine, serine, nonylamine, and a mixture of all four) were used to fertilize the soils in CF and BF every month for 1 year. Soil samples were collected every 2 months. Subsequently, soil microbial biomass and EEA were assayed. Results showed that the microbial biomass and EEA of soils fertilized with urea and amino acids increased significantly, whereas those fertilized with nonylamine and mixed N decreased significantly. Urea and amino acid fertilizations had a more positive influence on EEA of BF than on those of CF. Nonylamine fertilization had a more negative influence on EEA of CF than on those of BF. Organic N fertilization shifted soil microbial biomass away from the excretion of N-degrading enzymes and toward the excretion of C-degrading enzymes. These results suggest that organic N type is an important factor that affects soil microbial biomass, EEA, and their relationship. Organic N deposition may seriously affect soil C and N cycling, as well as carbon dioxide releasing from the soils by influencing microbial activities and biomass. This study thereby provides evidence that soil microorganisms have strong feedback to different forms of organic N deposition.     

The effect of nutrient deposition on bacterial communities in Arctic tundra soil

The microbial communities of high‐latitude ecosystems are expected to experience rapid changes over the next century due to climate warming and increased deposition of reactive nitrogen, changes that will likely affect microbial community structure and function. In moist acidic tundra (MAT) soils on the North Slope of the Brooks Range, Alaska, substantial losses of C and N were previously observed after long‐term nutrient additions. To analyse the role of microbial communities in these losses, we utilized 16S rRNA gene tag pyrosequencing coupled with community‐level physiological profiling to describe changes in MAT bacterial communities after short‐ and long‐term nutrient fertilization in four sets of paired control and fertilized MAT soil samples. Bacterial diversity was lower in long‐term fertilized plots. The Acidobacteria were one of the most abundant phyla in all soils and distinct differences were noted in the distributions of Acidobacteria subgroups between mineral and organic soil layers that were also affected by fertilization. In addition, Alpha‐ and Gammaproteobacteria were more abundant in long‐term fertilized samples compared with control soils. The dramatic increase in sequences within the Gammaproteobacteria identified as Dyella spp. (order Xanthomonadales) in the long‐term fertilized samples was confirmed by quantitative PCR (qPCR) in several samples. Long‐term fertilization was also correlated with shifts in the utilization of specific substrates by microbes present in the soils. The combined data indicate that long‐term fertilization resulted in a significant change in microbial community structure and function linked to changes in carbon and nitrogen availability and shifts in above‐ground plant communities.     

Experimental litterfall manipulation drives large and rapid changes in soil carbon cycling in a wet tropical forest

Global changes such as variations in plant net primary production are likely to drive shifts in leaf litterfall inputs to forest soils, but the effects of such changes on soil carbon (C) cycling and storage remain largely unknown, especially in C‐rich tropical forest ecosystems. We initiated a leaf litterfall manipulation experiment in a tropical rain forest in Costa Rica to test the sensitivity of surface soil C pools and fluxes to different litter inputs. After only 2 years of treatment, doubling litterfall inputs increased surface soil C concentrations by 31%, removing litter from the forest floor drove a 26% reduction over the same time period, and these changes in soil C concentrations were associated with variations in dissolved organic matter fluxes, fine root biomass, microbial biomass, soil moisture, and nutrient fluxes. However, the litter manipulations had only small effects on soil organic C (SOC) chemistry, suggesting that changes in C cycling, nutrient cycling, and microbial processes in response to litter manipulation reflect shifts in the quantity rather than quality of SOC. The manipulation also affected soil CO ₂ fluxes; the relative decline in CO ₂ production was greater in the litter removal plots (?22%) than the increase in the litter addition plots (+15%). Our analysis showed that variations in CO ₂ fluxes were strongly correlated with microbial biomass pools, soil C and nitrogen (N) pools, soil inorganic P fluxes, dissolved organic C fluxes, and fine root biomass. Together, our data suggest that shifts in leaf litter inputs in response to localized human disturbances and global environmental change could have rapid and important consequences for belowground C storage and fluxes in tropical rain forests, and highlight differences between tropical and temperate ecosystems, where belowground C cycling responses to changes in litterfall are generally slower and more subtle.     

森林生态系统碳循环对全球氮沉降的响应

森林土壤和植被储存着全球陆地生态系统大约46%的碳,在全球碳平衡中起着非常重要的作用。过去几十年来,森林生态系统的碳循环和碳吸存受到了全球氮沉降的深刻影响,因为氮沉降改变了陆地生态系统的生产力和生物量积累。以欧洲和北美温带森林区域开展的研究为基础,综述了氮沉降对植物光合作用、土壤呼吸、土壤DOM及林木生长的影响特征和机理,探讨了森林生态系统碳动态对氮沉降响应的不确定性因素。热带森林C、N循环与大部分温带森林不同,人为输入的氮对热带生态系统过程的影响也可能不同,因此指出了在热带地区开展碳氮循环耦合研究的必要性和紧迫性。     

Long‐term nitrogen addition modifies microbial composition and functions for slow carbon cycling and increased sequestration in tropical forest soil

Nitrogen (N) deposition is a component of global change that has considerable impact on belowground carbon (C) dynamics. Plant growth stimulation and alterations of fungal community composition and functions are the main mechanisms driving soil C gains following N deposition in N‐limited temperate forests. In N‐rich tropical forests, however, N deposition generally has minor effects on plant growth; consequently, C storage in soil may strongly depend on the microbial processes that drive litter and soil organic matter decomposition. Here, we investigated how microbial functions in old‐growth tropical forest soil responded to 13 years of N addition at four rates: 0 (Control), 50 (Low‐N), 100 (Medium‐N), and 150 (High‐N) kg N ha^?1 year^?1. Soil organic carbon (SOC) content increased under High‐N, corresponding to a 33% decrease in CO₂ efflux, and reductions in relative abundances of bacteria as well as genes responsible for cellulose and chitin degradation. A 113% increase in N₂O emission was positively correlated with soil acidification and an increase in the relative abundances of denitrification genes (narG and norB). Soil acidification induced by N addition decreased available P concentrations, and was associated with reductions in the relative abundance of phytase. The decreased relative abundance of bacteria and key functional gene groups for C degradation were related to slower SOC decomposition, indicating the key mechanisms driving SOC accumulation in the tropical forest soil subjected to High‐N addition. However, changes in microbial functional groups associated with N and P cycling led to coincidentally large increases in N₂O emissions, and exacerbated soil P deficiency. These two factors partially offset the perceived beneficial effects of N addition on SOC storage in tropical forest soils. These findings suggest a potential to incorporate microbial community and functions into Earth system models considering their effects on greenhouse gas emission, biogeochemical processes, and biodiversity of tropical ecosystems.     

Interactive effects of warming and increased nitrogen deposition on 15N tracer retention in a temperate old field: seasonal trends

The extent to which increased atmospheric nitrogen (N) deposition will drive changes in plant productivity and species composition over the next century will depend on how other influential global change factors, such as climate warming, affect the N retention of ecosystems. We examined the interactive effects of simulated climate warming and N deposition on the recoveries of ¹⁵N‐labeled ammonium and ¹⁵N‐labeled nitrate tracers added as a pulse to grass‐dominated, temperate old‐field plots at spring thaw. In addition to the year‐round warming treatment, a winter‐only warming treatment was applied to a set of plots to explore the contribution of this component of climate warming to the overall warming effect. By the end of the plant growing season, there was approximately twice as much ¹⁵N enrichment in the plant roots and bulk soil from ¹⁵NH₄⁺‐addition plots than from ¹⁵NO₃^?‐addition plots, but there were no effects of warming or N fertilization on ¹⁵N recovery. Over winter, approximately half of the excess ¹⁵N present in plant shoots was lost, which corresponded with large ¹⁵N losses from bulk soil in N fertilized plots and large ¹⁵N increases in bulk soil in nonfertilized plots. By the next spring, there was decreased ¹⁵N recovery in plants in response to N fertilization, which was largely offset by increases in plant ¹⁵N recovery in response to year‐round warming. However, ¹⁵N retention in bulk soil, where the major part of the ¹⁵N label was recovered, was approximately 40% higher in nonfertilized plots than in N fertilized plots. Overall, our results indicate that climate warming increases plant N sequestration in this system but this effect is overwhelmed by the overall effect of nitrogen deposition on ecosystem N losses.     

Mechanisms underlying limited soil carbon gains in perennial and cover‐cropped bioenergy systems revealed by stable isotopes

Removal of biomass for bioenergy production may decrease soil organic carbon. While perennials or cover‐cropped grains often have greater root production than annual grain crops, they variably impact soil carbon and underlying mechanisms remain unclear. We used high‐frequency measurements of soil respiration and natural abundance carbon stable isotopes to differentiate respiration sources, pool sizes, and decomposition rate constants during a 10 month incubation of soils collected to 1 m depth from a 10 year old field experiment in Iowa, United States. Conversion of corn–soybean rotations to reconstructed prairies or addition of a rye cover crop to continuous corn significantly altered respiration sources and dynamics of fast‐ and slow‐cycling carbon (turnover times of weeks to months–years, respectively), but had little effect on bulk soil carbon and several extractable pools (except in fertilized prairie). Both unfertilized and fertilized prairies increased slow‐cycling carbon pools relative to annual crops, but only in 0–25 cm soil. Compared with fertilized prairie, the unfertilized prairie significantly increased decomposition rates of fast‐ and slow‐cycling carbon pools in 0–25 cm soil, likely explaining the lack of significant bulk soil carbon accrual despite twofold greater root production. Carbon derived from C₄ plants decomposed faster than C₃‐derived carbon across all depths and cropping systems and contributions of C₃‐carbon to respiration increased with depth. Respiration of cover crop‐derived carbon was greatest in 0–25 cm soil but comprised >25% of respiration below 25 cm, implying a disproportionate impact of the cover crop on deep soil metabolism. However, the cover crop also increased the decomposition rates of fast‐ and slow‐cycling carbon pools and decreased their pool sizes across all depths relative to corn without a cover crop. Despite their notable environmental benefits, neither unfertilized perennials nor cover crops necessarily promote rapid soil carbon sequestration relative to conventional annual bioenergy systems because of concomitant increases in decomposition.     

Biodiversity,Nitrogen Deposition,and CO<Subscript>2</Subscript> Affect Grassland Soil Carbon Cycling but not Storage

Grasslands are globally widespread and capable of storing large amounts of carbon (C) in soils, and are generally experiencing increasing atmospheric CO₂, nitrogen (N) deposition, and biodiversity losses. To better understand whether grasslands will act as C sources or sinks in the future we measured microbial respiration in long-term laboratory incubations of soils collected from a grassland field experiment after 9 years of factorial treatment of atmospheric CO₂, N deposition, and plant species richness on a deep and uniformly sandy soil. We fit microbial soil respiration rates to three-pool models of soil C cycling to separate treatment effects on decomposition and pool sizes of fast, slow, and resistant C pools. Elevated CO₂ decreased the mean residence time (MRT) of slow C pools without affecting their pool size. Decreasing diversity reduced the size and MRT of fast C pools (comparing monocultures to plots planted with 16 species), but increased the slow pool MRT. N additions increased the size of the resistant pool. These effects of CO₂, N, and species-richness treatments were largely due to plant biomass differences between the treatments. We found no significant interactions among treatments. These results suggest that C sequestration in sandy grassland soils may not be strongly influenced by elevated CO₂ or species losses. However, high N deposition may increase the amount of resistant C in these grasslands, which could contribute to increased C sequestration.     

Effects of plant diversity,N fertilization,and elevated carbon dioxide on grassland soil N cycling in a long‐term experiment

The effects of global environmental changes on soil nitrogen (N) pools and fluxes have consequences for ecosystem functions such as plant productivity and N retention. In a 13‐year grassland experiment, we evaluated how elevated atmospheric carbon dioxide (CO₂), N fertilization, and plant species richness alter soil N cycling. We focused on soil inorganic N pools, including ammonium and nitrate, and two N fluxes, net N mineralization and net nitrification. In contrast with existing hypotheses, such as progressive N limitation, and with observations from other, often shorter, studies, elevated CO₂ had relatively static and small, or insignificant, effects on soil inorganic N pools and fluxes. Nitrogen fertilization had inconsistent effects on soil N transformations, but increased soil nitrate and ammonium concentrations. Plant species richness had increasingly positive effects on soil N transformations over time, likely because in diverse subplots the concentrations of N in roots increased over time. Species richness also had increasingly positive effects on concentrations of ammonium in soil, perhaps because more carbon accumulated in soils of diverse subplots, providing exchange sites for ammonium. By contrast, subplots planted with 16 species had lower soil nitrate concentrations than less diverse subplots, especially when fertilized, probably due to greater N uptake capacity of subplots with 16 species. Monocultures of different plant functional types had distinct effects on N transformations and nitrate concentrations, such that not all monocultures differed from diverse subplots in the same manner. The first few years of data would not have adequately forecast the effects of N fertilization and diversity on soil N cycling in later years; therefore, the dearth of long‐term manipulations of plant species richness and N inputs is a hindrance to forecasting the state of the soil N cycle and ecosystem functions in extant plant communities.     

Nonlinear root-derived carbon sequestration across a gradient of nitrogen and phosphorous deposition in experimental mesocosms

Enhanced sequestration of plant‐carbon (C) inputs to soil may mitigate rising atmospheric carbon dioxide (CO₂) concentrations and related climate change but how this sequestration will respond to anthropogenic nitrogen (N) and phosphorous (P) deposition is uncertain. We couple isotope, soil C fractionation and mesocosm techniques to assess the sequestration of plant‐C inputs, and their partitioning into C pools with different sink potentials, under an experimental gradient of N and P deposition (0, 10, 30, 60 and 100 kg N ha^?1 yr^?1; and 0, 2, 6, 12 and 20 kg P ha^?1 yr^?1). We hypothesized that N deposition would increase sequestration, with the majority of the C being sequestered in faster cycling soil pools because N deposition has been shown to accelerate the turnover of these pools while decelerating the turnover of slower cycling pools. In contrast to this hypothesis, sequestration into all soil C pools peaked at intermediate levels of N deposition. Given that P amendment has been shown to cause a net loss of soil C, we postulated that P deposition would decrease sequestration. This expectation was not supported by our data, with sequestration generally being greater under P deposition. When soils were amended simultaneously with N and P, neither the shape of the sequestration relationship across the deposition gradient, nor the observed sequestration at the majority of the deposition rates, was statistically predictable from the effects of N and P in isolation. The profound nonlinearities we observed, both for total sequestration responses and the partitioning of C into soil pools with different sink potentials, suggests that the rates of N and P deposition to ecosystems will be the critical determinant of whether they enhance or decrease the long‐term sequestration of fresh plant‐C inputs to soils.     

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.     

Microbial responses to warming enhance soil carbon loss following translocation across a tropical forest elevation gradient

Tropical soils contain huge carbon stocks, which climate warming is projected to reduce by stimulating organic matter decomposition, creating a positive feedback that will promote further warming. Models predict that the loss of carbon from warming soils will be mediated by microbial physiology, but no empirical data are available on the response of soil carbon and microbial physiology to warming in tropical forests, which dominate the terrestrial carbon cycle. Here we show that warming caused a considerable loss of soil carbon that was enhanced by associated changes in microbial physiology. By translocating soils across a 3000 m elevation gradient in tropical forest, equivalent to a temperature change of ± 15 °C, we found that soil carbon declined over 5 years by 4% in response to each 1 °C increase in temperature. The total loss of carbon was related to its original quantity and lability, and was enhanced by changes in microbial physiology including increased microbial carbon‐use‐efficiency, shifts in community composition towards microbial taxa associated with warmer temperatures, and increased activity of hydrolytic enzymes. These findings suggest that microbial feedbacks will cause considerable loss of carbon from tropical forest soils in response to predicted climatic warming this century.