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The ratio of β-1,4-glucosidase (BG) to β-1,4-N-acetylglucosaminidase (NAG) activity (BG:NAG ratio) is often used as an indicator of the relative resource allocation of soil microbes to C acquisition compared with N (the ecoenzymatic stoichiometry hypothesis). An increasing number of recent studies have used this index to assess the nutrient status of microbes. However, the validity of this index for assessing the nutrient status of microbes is not well tested. In this study, we collected published data and tested that validity by investigating whether N fertilization elevated the BG:NAG ratio, to test the assumption derived from the ecoenzymatic stoichiometry hypothesis that microbes reduce their allocation to the N-acquiring enzyme (NAG) under N-enriched conditions. Of the observations, 54% (82/151) did not support the hypothesis because those studies showed lower BG:NAG ratios in N-enriched soils than under ambient conditions (i.e., N was not enriched), especially when the ambient BG:NAG ratio was higher than 2.0 (77%, 59/77). This suggests that the BG:NAG ratio does not necessarily indicate the microbial status for C or N limitation. Rather, we hypothesized that NAG targets chitin or peptidoglycan, which accumulate at later stages of decomposition and that the variation in BG:NAG explains the decomposition stage. A negative correlation of BG:NAG ratio with polyphenol oxidase activity, which generally increases with decomposition, supported our hypothesis.

Asymbiotic nitrogen (N) fixation is an important source of new N in ecosystems, and is sensitive to atmospheric N deposition. However, there is limited understanding of asymbiotic N fixation and its response to N deposition in the context of forest rehabilitation. In this study, we measured N fixation rates (acetylene reduction) in different ecosystem compartments (i.e. soil, forest floor, moss Syrrhopodon armatus, and canopy leaves) in a disturbed and a rehabilitated subtropical forest in southern China, under 12 years of N treatments: control, low N addition (50 kg N ha− 1 yr− 1), and medium N addition (100 kg N ha− 1 yr− 1). The rehabilitated forest had higher nutrient (e.g. N) availability than the disturbed forest. In control plots, N fixation rates in forest floor were higher in the rehabilitated forest than in the disturbed forest, but N fixation rates in other compartments (soil, S. armatus, and canopy leaves) were comparable between the forests. Nitrogen addition significantly suppressed N fixation in soil, forest floor, S. armatus, and canopy leaves in the disturbed forest, but had no significant effect on those compartments in the rehabilitated forest. The main reasons for the negative effects of N addition on N fixation in the disturbed forest were NH4+ inhibition (soil), the P and C limitation (forest floor), and the reduced N dependence on canopy N-fixers (S. armatus and canopy leaves). We conclude that asymbiotic N fixation does not decline with increasing N availability after rehabilitation in the study forests. The inhibitory effects of N addition on asymbiotic N fixation occurred in the disturbed forest but not in the rehabilitated forest, indicating that forest rehabilitation may change the response of ecosystem function (i.e. N fixation) to N deposition, which merits further study in other tropical and subtropical regions.

Reforested ecosystems are becoming a dominant forest cover type in the tropics where anthropogenic nitrogen (N) deposition has been increasing greatly. Although understory plants play an important role in ecosystem structure and functioning, it remains unclear how long-term N deposition affects eco-physiological traits of understory species. To address this question, we expanded a ten-year simulated N deposition experiment in a tropical reforested ecosystem, with variable N addition levels as Control (no addition, N0), 50 kg N ha −1 yr −1 (N50), and 100 kg N ha −1 yr −1 (N100). Leaf physiological traits associated with photosynthetic capacity, foliar elements and N storage of seven dominant understory species were measured. Results showed that both medium-light species and shade-tolerant species had little physiological responses to N addition. However, one high-light species ( Alchornea trewioides ) responded significantly to N addition, showing decreases foliar N and Pmax. We further found that Pmax of Alchornea trewioides was negatively correlated with the canopy closure which increased under long-term N addition. Our findings suggest that understory plant growth may be more limited by light rather than N availability, particularly under chronic high N deposition; and N deposition can suppress understory high-light species through accelerating canopy closure in some tropical reforested ecosystems.

Natural abundance of 15N (δ15N) in plants and soils can provide time-integrated information related to nitrogen (N) cycling within ecosystems, but it has not been well tested in warm and humid subtropical forests. In this study, we used ecosystem δ15N to assess effects of increased N deposition on N cycling in an old-growth broad-leaved forest and a secondary pine forest in a high-N-deposition area in southern China. We measured δ15N of inorganic N in input and output fluxes under ambient N deposition, and we measured N concentration (%N) and δ15N of major ecosystem compartments under ambient deposition and after decadal N addition at 50 kg N ha−1yr−1, which has a δ15N of −0.7 ‰. Our results showed that the total inorganic N in deposition was 15N-depleted (−10 ‰) mainly due to high input of strongly 15N-depleted NH4+-N. Plant leaves in both forests were also 15N-depleted (−4 to −6 ‰). The broad-leaved forest had higher plant and soil %N and was more 15N-enriched in most ecosystem compartments relative to the pine forest. Nitrogen addition did not significantly affect %N in the broad-leaved forest, indicating that the ecosystem pools are already N-rich. However, %N was marginally increased in pine leaves and significantly increased in understory vegetation in the pine forest. Soil δ15N was not changed significantly by the N addition in either forest. However, the N addition significantly increased the δ15N of plants toward the 15N signature of the added N, indicating incorporation of added N into plants. Thus, plant δ15N was more sensitive to ecosystem N input manipulation than %N in these subtropical forests. We interpret the depleted δ15N of plants as an imprint from the high and 15N-depleted N deposition that may dominate the effects of fractionation that are observed in most warm and humid forests. Fractionation during the steps of N cycling could explain the difference between negative δ15N in plants and positive δ15N in soils, and the increase in soil δ15N with depths. Nevertheless, interpretation of ecosystem δ15N from high-N-deposition regions needs to include data on the deposition 15N signal.

Phosphorus (P) availability can affect nitrogen (N) dynamics in forest soil, and this effect might depend largely on the soil N status of forest ecosystems. So far, however, this view has not been well tested among forests with contrasting N status. Here, we used a 6-year experiment with additions of N and P to evaluate the effects of P availability and its interaction with N availability on soil N dynamics in one N-saturated and two N-limited tropical forests in southern China. Soil inorganic N concentrations and rates of N mineralization, nitrification, nitrous oxide (N2O) emission and nitrate leaching were measured. Our results showed that addition of P alone changed soil N dynamics in the N-saturated forest only; it accelerated rates of soil N transformation and decreased rates of N2O emission and nitrate leaching, but had no significant effects on N dynamics in the two N-limited forests. Furthermore, compared with the addition of N alone, addition of both N and P caused significant increases in the rates of net N mineralization and nitrification and a significant decrease in N2O emission in the two N-limited forests. Our results suggest that P availability stimulates soil N dynamics only when the ecosystem is saturated with N or there is considerable N deposition. Highlights We compared the effects of P addition on soil N dynamics among forests with different N status. Addition of P alone changed N dynamics in the N-saturated forest, but not in N-limited forests. Combined N and P additions had a larger effect on N dynamics than N addition alone. Phosphorus addition affects N dynamics only when an ecosystem has considerable N status.

Developing an understanding of the response of soil organic carbon (SOC) to N addition is critical to quantify and predict the terrestrial carbon uptake under increasing N deposition in the future. However, results from field studies on the response of SOC content and composition to N addition are highly variable across different ecosystems. The interpretation of SOC responses to N addition are often complicated by the differences in climate, soil substrate and other factors. To address this question, we measured SOC and its components in adjacent broadleaved and coniferous subtropical forests after 14 years of N addition. SOC in the top 50 cm increased by 2.1 kg m−2, 1.8 kg m−2 and 1.2 kg m−2 for low, medium and high rates of N addition in the broadleaved forest, but did not change significantly in the coniferous forest. Increased SOC in the broadleaved forest was contributed by the significant increases in particulate organic carbon (POC), humus organic carbon (HOC) in the 0–10 cm and 30–50 cm soil layers and resistant organic carbon (ROC) in the 0–10 cm soil layer. 13C nuclear magnetic resonance (NMR) spectra of coarse SOC revealed a decrease in easily decomposed carbon (C) and a shift in recalcitrant C. The increased SOC accumulation in the broadleaved forest was largely driven by altered rates of organic matter decomposition, rather than C inputs to soil. Land-history and low nutrient availability may have contributed to the lack of significant impact of N addition on SOC in the coniferous forest. Our results suggested the different controls of SOC accumulation and less sensitivity of SOC chemical composition at the molecular level to N addition in the two subtropical forest soils.

The prevailing paradigm for soil microbial activity in tropical forests is that microbial activity is limited by phosphorus (P) availability. Thus, exogenous P addition should increase rates of organic matter decomposition. Studies have also confirmed that soil respiration is accelerated when P is added experimentally. However, we hypothesize that the increased rates of soil microbial respiration could be due to the release of organic material from the surface of soil minerals when P is added, because P is more successful at binding to soil particles than organic compounds. In this study, we demonstrate that P addition to soil is associated with significantly higher dissolved organic carbon (DOC) content in a tropical evergreen forest in southern China. Our results indicate that P fertilization stimulated soil respiration but suppressed litter decomposition. Results from a second sorption experiment revealed that the recovery ratio of added DOC in the soil of a plot fertilized with P for 9 years was larger than the ratio in the soil of a non-fertilized plot, although the difference was small. We also conducted a literature review on the effects of P fertilization on the decomposition rates of litter and soil organic matter at our study site. Previous studies have consistently reported that P addition led to higher response ratios of soil microbial respiration than litter decomposition. Therefore, experiments based on P addition cannot be used to test whether microbial activity is P-limited in tropical forest soils, because organic carbon desorption occurs when P is added. Our findings suggest that the prevailing paradigm on the relationship between P and microbial activity in tropical forest soils should be re-evaluated.

Tropical forests with highly-weathered soils are considered to be high sensitive to nitrogen (N) deposition and subsequent soil acidification. Phosphorus (P) shortage in tropical forest ecosystems is suggested to be one important factor that makes the ecosystems more vulnerable to N-derived acidification. However, it remains poorly understood on how changes in soil P availability affect soil acidification processes in humid tropical zones with elevated atmospheric N deposition. To address this question, we conducted a long-term N and P addition experiment in an N-rich tropical forest, consisting of four treatments: control, N-addition, P-addition, and NP-addition, respectively (both N and P are 150 kg ha − 1 yr − 1 ). We hypothesized that relieving P-limitation by exogenous P addition will mitigate N-derived soil acidification. Results showed that six-year N addition significantly decreased soil pH (by 0.23, p = 0.013), soil base saturation (BS) (by 32.8%, p = 0.0017), and increased the ratio of aluminum to calcium (Al/Ca) ( p = 0.015), suggesting that soil acidification was accelerated by N input. As we expected, P addition increased soil pH in the first few years, probably because of the increased biotic uptakes of nitrate and phosphate. However, after the first few years, continuous N addition promoted acidification and made the buffering effects by P addition invisible in our soils at the Al-buffering stage. Our results suggest that P inputs cannot alleviate excess N-derived soil acidification in N-rich tropical ecosystems, and that it is urgent to reduce reactive N emission for sustainable development of the affected ecosystems in the future.

Atmospheric nitrogen (N) deposition is generally thought to suppress soil methane (CH4) uptake in natural forests, and phosphorus (P) input may alleviate this negative effect. However, it remains unclear how N and P inputs control soil CH4 uptake in disturbed forests. In this study, soil CH4 uptake rates were measured in two disturbed forests, including a secondary forest (with previous, but not recent, disturbance) and a plantation forest (with recent continuous disturbance), in southern China for 34 months of N and/or P additions: control, N addition (150 kg N ha−1 yr−1), P addition (150 kg P ha−1 yr−1), and NP addition (150 kg N ha−1 yr−1 plus 150 kg P ha−1 yr−1). Mean CH4 uptake rate in control plots was significantly higher in the secondary forest (24.40 ± 0.81 µg CH4-C m−2 h−1) than in the plantation forest (17.07 ± 0.70 µg CH4-C m−2 h−1). CH4 uptake rate had negative relationships with soil water-filled pore space in both forests. In the secondary forest, N, P, and NP additions significantly decreased CH4 uptake by 39.7%, 27.8%, and 37.6%, respectively, but had no significant effects in the plantation forest, indicating that P input does not alleviate the suppression of CH4 uptake by N deposition. Taken together, our findings suggest that reducing anthropogenic disturbance, including harvesting of forest floor, and anthropogenic N and P inputs will increase soil CH4 uptake in disturbed forests, which is important in view of the increased trends in global warming during recent decades.

It is well documented that phosphorus (P) input stimulates biological nitrogen (N) fixation (BNF) in tropical forests with non-legume trees. However, in tropical legume forests with soil N enrichment and P deficiency, the effects of P availability and its combination with N on BNF remain poorly understood. In this study, we measured BNF rate in different compartments, i.e., bulk soil, forest floor, rhizosphere, and nodules, in two tropical plantations with legume trees Acacia auriculiformis (AA) versus non-legume trees Eucalyptus urophylla, (EU) in southern China after 4 years of P addition and combined N and P additions. The objective was to investigate how P addition and its combination with N addition regulate BNF in a tropical legume plantation, and to compare the effects with those in a non-legume plantation. Our results showed that total BNF rates were significantly higher in the P-addition plots than in the control plots by 27.4 ± 4.3 and 23.3 ± 1.7 % in the EU and AA plantations, respectively. Total BNF rates were significantly higher in the NP-addition plots than in the control plots by 27.7 ± 5.0 and 8.5 ± 1.4 % in the EU and AA plantations, respectively, which contrasted to our previous result that total BNF rates were significantly lower in N-addition plots than in the control plots in the AA plantation. These findings suggest that P input can stimulate BNF in tropical forest biome dominated by legume trees, even in consideration of elevated atmospheric N deposition. Thus, our study revealed the important role of P in regulating biological N input, which should be taken into account in the modeling of biogeochemical cycles in the future.

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 15 N labeling experiment performed in a N-rich old-growth tropical forest in southern China. We added 15 N tracer monthly as 15 NH415 NO3 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.

Rates of biological nitrogen fixation (BNF) were measured in ecosystem. In this study, we measured rates of BNF in ecosystem compartments (bulk soil, forest floor, rhizosphere soil, and nodule) in two mature tropical plantations in southern China with legume trees (Acacia auriculiformis, AA) and with non-legume trees (Eucalyptus urophylla, EU) after 4 years of nitrogen (N) fertilization (0, 50, and 100 kg N ha−1 year−1). BNF rates of bulk soil were comparable between plantations, while rates of rhizosphere soil were significantly higher in the EU plantation and rates of forest floor were significantly higher in the AA plantation. Thus, total BNF rates were comparable between plantations (AA = 6.04 kg N ha−1 year−1; EU= 6.42 kg N ha−1 year−1). In the AA plantation, N addition significantly decreased BNF rates in all measured compartments and thus the total rates. In the EU plantation, N addition did not change BNF rates of forest floor, but significantly decreased rates of bulk soil and increased rates of rhizosphere soil; thus, total rates did not change. Our findings provide evidence that forest type is an important factor regulating the effects of external N input on BNF, and suggest that elevated atmospheric N deposition in recent decades will suppress total N fixation in mature forests with legume trees but not in those with non-legume trees.

Dissolved organic matter (DOM) drives many fundamental biogeochemical processes (e.g., carbon storage, nutrient cycling, and soil development) in forest soil. However, the molecular-level characteristics of DOM derived from different types of tropical forest soils are poorly understood. Here, water samples at different soil depths (0, 20, and 40 cm) from tropical legume (Acacia auriculiformis, AA) and non-legume (Eucalyptus urophylla, EU) tree plantations were analyzed using absorption and fluorescence spectroscopy, Fourier transform ion cyclotron resonance mass spectrometry (FT-ICR MS), and solution-state 1H nuclear magnetic resonance (NMR) spectroscopy. The FT-ICR MS results indicated that DOM persisted in the soil, but its molecular composition notably shifted from low-mass (150–300 Da) and more-aromatic molecules to middle- (300–450 Da) and high-mass (>450 Da) and less-aromatic molecules with increasing soil depth. This was primarily mediated by consumption and mineral sorption of low-mass plant-derived DOM (e.g., low-mass carbohydrates and polyphenols) and further formation of larger microbial products (e.g., protein-like and lipid-like compounds). In addition, a higher abundance of microbial-derived molecules (e.g., protein-like and carboxyl-rich alicyclic molecules) was found at the legume plantation relative to the non-legume plantation, which suggests a faster microbial turnover of DOM. Also, the legume plantation had greater enrichment of middle- and high-mass and condensed aromatic-like DOM components in soils. These findings improve our understanding of the drivers that mediate the response of DOM to soil depth and tree species in tropical plantations.

Urbanization, the migration of populations from rural to urban areas, has been causing great stress on natural environments, leading to air pollution and nitrogen (N) deposition, negatively affecting forest health. Although there is evidence that urbanization has changed forest N cycling, little is known about whether urbanization also changes the availability of phosphorus (P), which is important for plant growth and forest productivity. To address this question, we carried out a survey in the Pearl River Delta region, the world's largest urban area in southern China, using two types of representative forests, the evergreen broadleaf forests (BFs) and pine plantations (PPs). The leaf N:P ratios in the two forest types were high (20-50) with a significant increasing pattern along the rural-to-urban gradient. The ratios of leaf P:K and P:Na declined along the rural-to-urban gradient, whereas leaf P content did not change in BF but decreased in PP along the rural-to-urban gradient, suggesting that leaf P became limiting along urbanization. The abundance of actinomycetes and gram-negative bacteria decreased along the rural-to-urban gradient, indicating the negative effects of urbanization on soil microorganisms. Principal component analysis indicated that divergent key factors respond to the urbanization and affect plant P limitation in the two forest types. In BF, broadleaf trees showed a greater response to N deposition from urbanization indicating direct leaf N uptake from N deposition is a key factor for plant P limitation. Alternatively, in PP, our findings suggest soil acidification is an important factor accelerating plant P limitation. Our study revealed that urbanization intensifies plant P limitation in subtropical forests, and the effects vary depending on forest types. Our findings provide empirical information to support the management of forest ecosystems and evaluation of urbanization effects on forest health.

Sulfur dioxide emissions have been regulated at a global scale; sulfur (S) deposition no longer contributes to soil acidification instead of an alleviation effect in temperate regions; however, it remains unclear whether S deposition still contributes to soil acidification in the tropics. The Pearl River Delta (PRD), South China, has been suffering serious soil acidification, but the contribution of S deposition was ignored because of the regulation of S emission since 2001. Here, we chose the evergreen broadleaf forests, which are the typical forest type at the regional scale in PRD to examine the contribution of S deposition and its characteristics in this acidification, based on an established urban–rural gradient in the range of 260 km. A substantial acidification was evidenced by the significant decline of soil pH from rural to urban sites, with mean pH values decreased by more than 0.60 U through the whole 40-cm depths. However, there was no significant difference in soil pH from 0–10 cm, 10–20 cm, and to 20–40 cm at each site (P > 0.05). Acid-neutralizing capacity (ANC) showed a similar trend to soil pH, with a significant decline along the urbanization gradient and no significant effect of soil depths. Soil sulfate (SO42−), as the most abundant species in ANC, contributed greatly to soil acidification for the whole 40-cm depth, as shown by the significant positive relationships between it with soil pH and base cations. Soils also exhibited the depletion of base cations with low base saturation (

Anthropogenic nitrogen (N) deposition has accelerated terrestrial N cycling at regional and global scales, causing nutrient imbalance in many natural and seminatural ecosystems. How added N affects ecosystems where N is already abundant, and how plants acclimate to chronic N deposition in such circumstances, remains poorly understood. Here, we conducted an experiment employing a decade of N additions to examine ecosystem responses and plant acclimation to added N in an N-rich tropical forest. We found that N additions accelerated soil acidification and reduced biologically available cations (especially Ca and Mg) in soils, but plants maintained foliar nutrient supply at least in part by increasing transpiration while decreasing soil water leaching below the rooting zone. We suggest a hypothesis that cation-deficient plants can adjust to elevated N deposition by increasing transpiration and thereby maintaining nutrient balance. This result suggests that long-term elevated N deposition can alter hydrological cycling in N-rich forest ecosystems.

Nitrogen (N) deposition in China has increased greatly, but the general impact of elevated N deposition on carbon (C) dynamics in Chinese terrestrial ecosystems is not well documented. In this study we used a meta-analysis method to compile 88 studies on the effects of N deposition C cycling on Chinese terrestrial ecosystems. Our results showed that N addition did not change soil C pools but increased above-ground plant C pool. A large decrease in below-ground plant C pool was observed. Our result also showed that the impacts of N addition on ecosystem C dynamics depend on ecosystem type and rate of N addition. Overall, our findings suggest that 1) decreased below-ground plant C pool may limit long-term soil C sequestration; and 2) it is better to treat N-rich and N-limited ecosystems differently in modeling effects of N deposition on ecosystem C cycle.

Atmospheric nitrogen (N) deposition has elevated rapidly in tropical regions where N2-fixing tree species are widespread. However, the effect of N deposition on litter decomposition in forests with N2-fixing tree species remains unclear. We examined the effect of N addition on litter decomposition and nutrient release in two tropical plantations with Acacia auriculiformis (AA, N2-fixing) and Eucalyptus urophylla (EU, non-N2-fixing) in South China. Three levels of N additions were conducted: control, medium-N (50 kg N ha−1 yr.−1) and high-N (100 kg N ha−1 yr.−1) in each plantation. Initial decomposition rate (k a ) for the control plots was faster in the AA plantation than in the EU plantation, but later in decomposition, larger fraction of slowly decomposing litter (A) remained in the former. N addition increased the slow fraction (A), decreasing soil microbial biomass and reducing acid-unhydrolyzable residue (AUR) degradation in the AA plantation. In the EU plantation, however, N additions significantly increased initial decomposition rate (k a ) and soil N availability. Furthermore, N addition decreased litter carbon and N release (in the AA plantation), while litter phosphorus release also decreased in both plantations. With ongoing N deposition in future, tropical plantations with N2-fixing tree species would potentially increase carbon accumulation and nutrient retention in forest floor by slowing litter decomposition.

Summary Nitrogen (N) saturation hypothesis suggests that when an ecosystem reaches N-saturation, continued N input will cause increased N leaching, nitrous oxide (N2O) emission, and N mineralization and nitrification rates. It also suggests that a different element will become the main limiting factor when N saturation has been reached. Although this hypothesis has been tested in temperate forests, whether they can be directly applied to N-saturated tropical forests remain poorly addressed. To test this hypothesis, soil inorganic N, soil N mineralization and nitrification rate, soil N2O emission rate and nitrate (NO3−) leaching rate were measured in an N-saturated old-growth tropical forest in southern China, after 6 years of N and P addition. We hypothesized that N addition would stimulate further N saturation, but P addition might alleviate N saturation. As expected, our results showed that six continuous years of experimental N addition did cause further N saturation, which was indicated by significant increases in soil inorganic N concentration, N2O emission and nitrate (NO3−) leaching. However, in contrast to our expectations, N addition significantly decreased in situ rates of net N mineralization and nitrification, which could be related to associated changes in enzyme activity and microbial community composition. On the other hand, P addition mitigated N saturation, as expected. Soil inorganic N concentration, N2O emission and NO3− leaching decreased significantly after P addition, but the net rates of N mineralization and nitrification were significantly increased. Our results provide a new understanding of the N saturation hypothesis, suggesting that the effects of long-term N deposition on net N mineralization and nitrification rates in N-saturated tropical forests can be negative and that P addition can alleviate N saturation in such tropical systems.

Atmospheric nitrogen (N) deposition has accelerated in the last several decades due to anthropogenic activities, such as nitrogen fertilization, N-fixing plants cultivation and fossil fuel and biomass combustion. Increasing N deposition has become one of the important factors regulating N cycle in forest ecosystems. Forest ecosystems can retain part of deposited N in soil by biotic and abiotic mechanisms, but when N inputs exceed the capacity of soil retention, N losses will aggravate in terms of N oxide emission and/or nitrate leaching. The excess N input has threatened ecosystem health via acidification and eutrophication, causing declines in terrestrial biodiversity and forest productivity in forest ecosystems of Europe and North America. Recently, China has become one of the three areas that undergo severe N deposition in the world. Impacts of N deposition on soil N cycle in Chinese forest ecosystems have received increasing concern. In this paper, we reviewed the processes of soil N cycle and their responses to atmospheric N deposition based on available literature. The objective is to enhance our understanding on how N deposition affects soil N cycle in forest ecosystems and provide scientific information for sustainable forest management. The review mainly includes the following four aspects: (1) processes of soil N cycle and their controlling factors. These processes include biological N fixation (BNF), decomposition, mineralization, nitrification, denitrification, N oxide emission and NO3−N leaching. The controlling factors of these processes are complicated and interactional. Only one of these factors altered may affect soil N cycle. For example, C/N is the factor that controls BNF, decomposition, mineralization and NO3−N leaching. (2) Research methods and current results about studies are related to the impact of N deposition on soil N cycle in forest ecosystems. In general, the research methods are long-term simulated N deposition experiment, N deposition gradient method, roof clean rain method and 15N tracing method. Effects of N deposition on soil N cycle vary depending on different initial N statuses and lengths of experiment. In “N-limited” forests, N deposition tended to have positive effect on soil N cycling processes, such as accelerating litter decomposition rate and N mineralization rate. However, such result generally showed in short-term fertilization experiments. In some long-term fertilization experiments, it showed that the negative effects would rise when the forests reached N saturation. Compared to “N-limited” forests in temperate region, N deposition tended to have negative or neutral effects in “N-rich” tropical forest. For example, N deposition promoted nitrification process in tropical forests. (3) Possible mechanisms for the effect of N deposition on soil N cycle: N deposition can affect soil N cycle through altering the chemical characteristic of forest substrates, the biomass and community composition of plant and microorganism. (4) Current problems and future research needs for the study about the effect of N deposition on soil N cycle: What role does regional diversity, changes in forest type, and interaction of carbon (C), N and phosphorus (P) play on the effect of N deposition on soil N cycle in forest ecosystems deserve our further study in the future.

Elevated nitrogen (N) deposition has dramatically altered soil phosphorus (P) and carbon (C) cycles in forests and altered extracellular enzyme activity. However, the effects of N addition and the interactive effects of combined N and P additions on soil enzyme (e.g. phosphatase and glucosidase) activity in different types of forests (young vs. old-growth) remain unclear. To better understand this, a long-term N–P fertilization experiment was initiated in January 2007 in two subtropical forests (an old-growth monsoon evergreen broadleaf forest (MEBF) and a young Masson pine forest (MPF)) in southern China. Four treatments were established, including control (no nutrient addition), N addition (150 kg N ha−1 yr−1), P addition (150 kg P ha−1 yr−1) and NP addition (150 kg N ha−1 yr−1 plus 150 kg P ha−1 yr−1). Soil physicochemical properties, acid phosphatase (APA-s) and β-glucosidase (BGA-s) activity per soil were measured in July 2012 and July 2013. Both APA-s and BGA-s were higher in MEBF than in MPF. N addition significantly stimulated APA-s in MEBF and BGA-s in MPF. P addition significantly suppressed APA-s in both forests and BGA-s in MEBF. Moreover, P addition decreased the stimulating effect of N addition on APA-s in MEBF and on BGA-s in MPF. Our results suggest that (1) N addition may exacerbate soil P limitation in old-growth forests and result in the deficiency of easily available C in young forests and (2) P addition may mitigate these negative effects of N addition in both forest types. Our findings suggest that P fertilization may be an effective practice to improve soil P availability in old-growth forests and soil C availability in young forests under N deposition condition.

Nitrogen (N) deposition has been shown to affect soil carbon (C) and N cycling in subtropical forests; however, the underlying microbial mechanisms are poorly understood. We used patterns of community-level physiological profiles and enzyme activities to assess the relative effects of the addition of four N levels (0, 50, 100, and 150 kg·ha−1·year−1) on the soil microbial community in three forest stands (pine, mixed, and broadleaf forests) in southern China, where the forests have been experimentally manipulated for over 8 years. In pine forests, N50 addition significantly increased microbial biomass carbon (MBC) concentration but decreased soil pH levels. N100 addition significantly increased soil peroxidase activity but decreased soil β-1,4-glucosidase activity. In broadleaf forests, N addition increased soil dissolved organic carbon (DOC) concentration and polyphenol oxidase activity but decreased soil MBC concentration and soil pH levels. N addition also significantly increased soil microbial metabolism activity (expressed as average well color development) in pine forest and broadleaf forest soils. However, the mixed forests responded slowly to N additions and exhibited no significant response of C-utilization profiles and soil enzyme activities. Principal component analysis of C-utilization data separated microbial communities with respect to N addition and forest successional stage. In addition, microbial C utilization was driven by soil pH levels. Although enzyme activities were correlated with soil MBC and microbial biomass nitrogen concentrations, stepwise regression results indicated that soil total carbon contents that were integrated with soil pH levels were key integrators of soil enzyme activities. Our results suggest that soil acidification due to N addition increased soil bacterial C utilization and enzyme activities.

Elevated anthropogenic nitrogen (N) deposition has become an important driver of soil acidification at both regional and global scales. It remains unclear, however, how long-term N deposition affects soil buffering capacity in tropical forest ecosystems and in ecosystems of contrasting land-use history. Here, we expand on a long-term N deposition experiment in three tropical forests that vary in land-use history (primary, secondary, and planted forests) in Southern China, with N addition as NH4NO3 of 0, 50, 100, and 150 kg N ha(-1) yr(-1), respectively. Results showed that all three forests were acid-sensitive ecosystems with poor soil buffering capacity, while the primary forest had higher base saturation and cation exchange capacity than others. However, long-term N addition significantly accelerated soil acidification and decreased soil buffering capacity in the primary forest, but not in the degraded secondary and planted forests. We suggest that ecosystem N status, influenced by different land-use history, is primarily responsible for these divergent responses. N-rich primary forests may be more sensitive to external N inputs than others with low N status, and should be given more attention under global changes in the future, because lack of nutrient cations is irreversible.

Understanding how phosphorus (P) and nitrogen (N) fertilization influence the dynamics of other nutrients [particularly sulfur (S)] is essential for improving the integrated nutrient management of tropical plantations. This study investigated how adding P and N fertilizers would affect S dynamics in the soils of two tropical forest plantations. Specifically, we tested whether the addition of P and N alters the availability of exchangeable S and microbial biomass S (MBS) and, hence, arylsulfatase activity (AS) and ratios of AS to C- and/or N-acquiring enzyme activity in the 0–5 cm soil layer were measured. The sites contained both control plots (no fertilizer application) and plots with 6-year continuous fertilizer (P and N) application. Soil exchangeable S clearly decreased, while AS activity marginally increased in P-fertilized plots. The ratios of soil AS activity to C- and/or N-acquiring enzyme activity tended to increase, possibly because the requirements of microbes for S relative to C and N were elevated, with P fertilization exacerbating shortages in S. In comparison, N fertilization did not influence exchangeable S, soil pH, and MBS. This lack of response might be attributed to continuous deposition of high levels of background N at the study sites. Soil AS activity and the ratios of soil AS activity to C- and/or N-acquiring enzyme activity tended to be lowered by N fertilization; thus, AS activity might be more sensitive to N fertilization than to soil pH. In conclusion, P fertilization must be carefully managed in tropical forest plantations to minimize its negative impact on soil S availability.

Differences in nitrogen (N) acquisition patterns between plant species are often reflected in the natural 15N isotope ratios (&delta, 15N) of the plant tissues, however, such differences are poorly understood for co-occurring plants in tropical and subtropical forests. To evaluate species variation in N acquisition traits, we measured leaf N concentration (%N) and &delta, 15N in tree and understory plant species under ambient N deposition (control) and after a decade of N addition at 50 kg N ha&minus, 1 yr&minus, 1 (N-plots) in an old-growth subtropical forest in southern China. We also measured changes in leaf &delta, 15N after one-year of 15N addition in both the control and N-plots. The results show consistent significant species variation in leaf %N in both control and N-plots, but decadal N addition did not significantly affect leaf %N. Leaf &delta, 15N values were also significantly different among the plant species both in tree and understory layers, and both in control and N-plots, suggesting differences in N acquisition strategies such as variation in N sources and dominant forms of N uptake and dependence on mycorrhizal associations among the co-occurring plant species. Significant differences between the plant species (in both control and N-plots) in changes in leaf &delta, 15N after 15N addition were observed only in the understory plants, indicating difference in access (or use) of deposited N among the plants. Decadal N addition had species-dependent effects on leaf &delta, 15N, suggesting the N acquisition patterns of these plant species are differently affected by N deposition. These results suggest that co-occurring plants in N-rich and subtropical forests vary in their N acquisition traits, these differences need to be accounted for when evaluating the impact of N deposition on N cycling in these ecosystems.

Natural abundance of 15N (δ15N) in plants and soils can provide integrated information on ecosystem nitrogen (N) cycling, but it has not been well tested in warm and humid sub-tropical forests. In this study, we examined the measurement of δ15N for its ability to assess changes in N cycling due to increased N deposition in an old-growth broadleaved forest and a secondary pine forest in a high N deposition area in southern China. We measured δ15N of inorganic N in input and output fluxes under ambient N deposition, and N concentration (N %) and δ15N of major ecosystem compartments under ambient and after decadal N addition at 50 kg N ha−1 yr−1. Our results showed that the N deposition was δ15N-depleted (−12 ‰) mainly due to high input of depleted NH4+-N. Plant leafs in both forest were also δ15N-depleted (−4 to −6 ‰). The old-growth forest had higher plant and soil N %, and was more 15N-enriched in most ecosystem compartments relative to the pine forest. Nitrogen addition did not significantly affect N % in both forests, indicating that the ecosystem pools are already N-rich. Soil δ15N was not changed significantly by the N addition in both forests. However, the N addition significantly increased the δ15N of plants toward the 15N signature of the added N (~ 0 ‰), indicating incorporation of added N into plants. Thus, plant δ15N was sensitive to ecosystem N input manipulation although N % was unchanged in these N-rich sub-tropical forests. We interpret the depleted δ15N values of plants as an imprint from the high and δ15N-depleted N deposition. The signal from the input (deposition or N addition) may override the enrichment effects of fractionation during the steps of N cycling that are observed in most warm and humid forests. Thus, interpretation of ecosystem δ15N values from high N deposition regions need to include data on the deposition δ15N signal.

Nitrogen (N) deposition is generally considered to increase soil nitrous oxide (N2O) emission in N-rich forests. In many tropical forests, however, elevated N deposition has caused soil N enrichment and further phosphorus (P) deficiency, and the interaction of N and P to control soil N2O emission remains poorly understood, particularly in forests with different soil N status. In this study, we examined the effects of N and P additions on soil N2O emission in an N-rich old-growth forest and two N-limited younger forests (a mixed and a pine forest) in southern China to test the following hypotheses: (1) soil N2O emission is the highest in old-growth forest due to the N-rich soil; (2) N addition increases N2O emission more in the old-growth forest than in the two younger forests; (3) P addition decreases N2O emission more in the old-growth forest than in the two younger forests; and (4) P addition alleviates the stimulation of N2O emission by N addition. The following four treatments were established in each forest: Control, N addition (150 kg N ha−1 yr−1), P addition (150 kg P ha−1 yr−1), and NP addition (150 kg N ha−1 yr−1 plus 150 kg P ha−1 yr−1). From February 2007 to October 2009, monthly quantification of soil N2O emission was performed using static chamber and gas chromatography techniques. Mean N2O emission was shown to be significantly higher in the old-growth forest (13.9 ± 0.7 µg N2O-N m−2 h−1) than in the mixed (9.9 ± 0.4 µg N2O-N m−2 h−1) or pine (10.8 ± 0.5 µg N2O-N m−2 h−1) forests, with no significant difference between the latter two. N addition significantly increased N2O emission in the old-growth forest but not in the two younger forests. However, both P and NP addition had no significant effect on N2O emission in all three forests, suggesting that P addition alleviated the stimulation of N2O emission by N addition in the old-growth forest. Although P fertilization may alleviate the stimulated effects of atmospheric N deposition on N2O emission in N-rich forests, this effect may only occur under high N deposition and/or long-term P addition, and we suggest future investigations to definitively assess this management strategy and the importance of P in regulating N cycles from regional to global scales.

The Effect of Simulated Nitrogen Deposition on the Emission of Carbonyl Compounds from Ormosia pinnata and Cinnamomum burmannii Elevated anthropogenic nitrogen (N) deposition has become a limiting factor for plants instead of a nutrient for its dramatic rising at globally scale, which greatly affects the process of carbon (C) cycle from individual plant to ecosystem, and even global levels. It could affect biogenic carbonyl emissions as an important component of carbon pool from plants, but this aspect has not yet been investigated. In this study, we performed a simulated N deposition experiment (100 kg N•ha-1•y-1) on two native tree saplings planted in a nursery in South China. Results showed that three main biogenic carbonyls, i.e. formaldehyde, acetaldehyde, and acetone from tree leaves all had distinct seasonal variations, with much higher emission rates in the wet season than in the dry season. Elevated N addition significantly depressed the emission of biogenic carbonyls in the dry season, but not in the wet season. No big differences were observed in carbonyls emissions from the two tree species responding to N deposition both in the wet season and in the dry season. Thus, we concluded that N deposition might drive plants to change carbon allocation and reduce carbon loss when as a limiting factor in the dry season. This finding is of significance for the theory of carbon allocation and plant adaptability strategy under elevated N deposition.

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

Soil acidification instead of alkalization has become a new environmental issue caused by urbanization. However, it remains unclear the characters and main contributors of this acidification. We investigated the effects of an urbanization gradient on soil acidity of Pinus massoniana forests in Pearl River Delta, South China. The soil pH of pine forests at 20-cm depth had significantly positive linear correlations with the distance from the urban core of Guangzhou. Soil pH reduced by 0.44 unit at the 0–10 cm layer in urbanized areas compared to that in non-urbanized areas. Nitrogen deposition, mean annual temperature and mean annual precipitation were key factors influencing soil acidification based on a principal component analysis. Nitrogen deposition showed significant linear relationships with soil pH at the 0–10 cm (for ammonium N ("Equation missing"-N), P -N), P -N, P

Increasing atmospheric nitrogen (N) deposition could profoundly impact community structure and ecosystem functions in forests. However, conventional experiments with understory addition of N (UAN) largely neglect canopy-associated biota and processes and therefore may not realistically simulate atmospheric N deposition to generate reliable impacts on forest ecosystems. Here we, for the first time, designed a novel experiment with canopy addition of N (CAN) vs. UAN and reviewed the merits and pitfalls of the two approaches. The following hypotheses will be tested: i) UAN overestimates the N addition effects on understory and soil processes but underestimates those on canopy-associated biota and processes, ii) with low-level N addition, CAN favors canopy tree species and canopy-dwelling biota and promotes the detritus food web and iii) with high-level N addition, CAN suppresses canopy tree species and other biota and favors rhizosphere food web. As a long-term comprehensive program, this experiment will provide opportunities for multidisciplinary collaborations, including biogeochemistry, microbiology, zoology and plant science to examine forest ecosystem responses to atmospheric N deposition.