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.     

氮沉降下不同碳添加模式对亚热带毛竹林土壤激发效应的影响

激发效应(PE)在调控陆地土壤碳(C)循环中发挥着重要作用,但在氮(N)沉降日益严重的亚热带森林生态系统中,不同C添加模式对PE的影响尚不清楚。本研究通过添加¹³C标记的葡萄糖,进行90 d的室内培养试验,探究不同施N水平下(0、20、80 kg N·hm^-2·a^-1)C添加模式(单次C添加、重复C添加)对土壤PE的影响。结果表明: 不同模式葡萄糖添加均显著增加了土壤有机C(SOC)矿化,产生了正PE,且单次的葡萄糖添加比重复添加引起的PE更大;随着施N水平的增加,PE显著减弱,表明N沉降抑制了毛竹林土壤激发。相关分析显示,累积PE与β-N-乙酰氨基葡萄糖苷酶(NAG)、过氧化物酶(PEO)活性呈显著负相关,与微生物生物量磷(MBP)、酸碱度(pH)呈显著正相关。综上,施N和C添加共同作用于土壤时,可以通过刺激亚热带森林中原生土壤有机质矿化而对土壤C储量产生强烈影响。本研究证明,单次C添加模式可能高估了外源易分解有机C对PE的影响,而忽略了N沉降对PE的影响,进而高估了森林SOC的矿化损失。     

Phosphorus Limitation of Microbial Processes in Moist Tropical Forests: Evidence from Short-term Laboratory Incubations and Field Studies

Although there is a widespread belief that phosphorus (P) limits basic ecosystem processes in moist tropical forests, direct tests of this supposition are rare. At the same time, it is generally believed that P does not limit soil microorganism respiration or growth in terrestrial ecosystems. We used natural gradients in P fertility created by soils of varying age underlying tropical rain forests in southwestern Costa Rica, combined with direct manipulations of carbon (C) and P supply, to test the effects of P availability on the decomposition of multiple forms of C, including dissolved organic carbon (DOC) and soil organic carbon (SOC). Results from a combination of laboratory and field experiments suggest that C decomposition in old, highly weathered oxisol soils is strongly constrained by P availability. In addition, P additions to these soils (no C added) also revealed that microbial utilization of at least labile fractions of SOC was also P limited. To our knowledge, this is the first direct evidence of P limitation of microbial processes in tropical rain forest soil. We suggest that P limitation of microbial decomposition may have profound implications for C cycling in moist tropical forests, including their potential response to increasing atmospheric carbon dioxide. Furthermore, this site is still relatively rich in P when compared to many other tropical forests on old soils; thus, we believe that P limitation of soil microorganisms throughout the humid tropics is a possibility.     

Soil C and N availability determine the priming effect: microbial N mining and stoichiometric decomposition theories

The increasing input of anthropogenically derived nitrogen (N) to ecosystems raises a crucial question: how does available N modify the decomposer community and thus affects the mineralization of soil organic matter (SOM). Moreover, N input modifies the priming effect (PE), that is, the effect of fresh organics on the microbial decomposition of SOM. We studied the interactive effects of C and N on SOM mineralization (by natural ¹³C labelling adding C₄‐sucrose or C₄‐maize straw to C₃‐soil) in relation to microbial growth kinetics and to the activities of five hydrolytic enzymes. This encompasses the groups of parameters governing two mechanisms of priming effects – microbial N mining and stoichiometric decomposition theories. In sole C treatments, positive PE was accompanied by a decrease in specific microbial growth rates, confirming a greater contribution of K‐strategists to the decomposition of native SOM. Sucrose addition with N significantly accelerated mineralization of native SOM, whereas mineral N added with plant residues accelerated decomposition of plant residues. This supports the microbial mining theory in terms of N limitation. Sucrose addition with N was accompanied by accelerated microbial growth, increased activities of β‐glucosidase and cellobiohydrolase, and decreased activities of xylanase and leucine amino peptidase. This indicated an increased contribution of r‐strategists to the PE and to decomposition of cellulose but the decreased hemicellulolytic and proteolytic activities. Thus, the acceleration of the C cycle was primed by exogenous organic C and was controlled by N. This confirms the stoichiometric decomposition theory. Both K‐ and r‐strategists were beneficial for priming effects, with an increasing contribution of K‐selected species under N limitation. Thus, the priming phenomenon described in ‘microbial N mining’ theory can be ascribed to K‐strategists. In contrast, ‘stoichiometric decomposition’ theory, that is, accelerated OM mineralization due to balanced microbial growth, is explained by domination of r‐strategists.     

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 suppression of in vitro growth of Pythium ultimum and disease incidence in relation to soil C and N availability

Experiments were designed to examine effects of the soil microbial community, C and N availability on in vitro growth of Pythium ultimum and its infection of cotton seedlings by manipulating the stage of cellulose decomposition, size and activity of microbial populations, and N availability. In comparison to the untreated control (CONT), cellulose addition alone (CELL) reduced soil nitrate by 35–80 fold, but had no significant effect on soil ammonium. Soil microbial biomass C (SMBC) increased over 2 fold in 14 days following cellulose addition, but significantly decreased in the following 10 days due to N limitation. Addition of both cellulose and N (NCELL) resulted in sustained SMBC for 24 days and significantly reduced in vitro P. ultimum growth and disease incidence. In vitro growth of P. ultimum and disease severity were consistently reduced in the order: CONT > CELL > NCELL. In vitro growth of P. ultimum was lower in soils previously incubated for 24 days than in those incubated for 14 days, and was most closely correlated to cumulative soil CO₂ evolution (CO2T). Correlations between P. ultimum growth rates and NO₃-N or total available N were substantial (p < 0.05), but much less significant than those between the growth rates and SMBC, microbial activity measured as CO₂ evolution rates or CO2T (p<0.0001). Addition of available N (NH₄NO₃) and C (glucose) just before the assays did not increase the in vitro growth of P. ultimum or disease severity on cotton seedlings, suggesting that time-dependent microbial processes or microbial metabolites significantly contributed to suppression of P. ultimum growth.     

The response of heterotrophic activity and carbon cycling to nitrogen additions and warming in two tropical soils

Nitrogen (N) deposition is projected to increase significantly in tropical regions in the coming decades, where changes in climate are also expected. Additional N and warming each have the potential to alter soil carbon (C) storage via changes in microbial activity and decomposition, but little is known about the combined effects of these global change factors in tropical ecosystems. In this study, we used controlled laboratory incubations of soils from a long‐term N fertilization experiment to explore the sensitivity of soil C to increased N in two N‐rich tropical forests. We found that fertilization corresponded to significant increases in bulk soil C concentrations, and decreases in C loss via heterotrophic respiration (P< 0.05). The increase in soil C was not uniform among C pools, however. The active soil C pool decomposed faster with fertilization, while slowly cycling C pools had longer turnover times. These changes in soil C cycling with N additions corresponded to the responses of two groups of microbial extracellular enzymes. Smaller active C pools corresponded to increased hydrolytic enzyme activities; longer turnover times of the slowly cycling C pool corresponded to reduced activity of oxidative enzymes, which degrade more complex C compounds, in fertilized soils. Warming increased soil respiration overall, and N fertilization significantly increased the temperature sensitivity of slowly cycling C pools in both forests. In the lower elevation forest, respired CO₂ from fertilized cores had significantly higher Δ¹⁴C values than control soils, indicating losses of relatively older soil C. These results indicate that soil C storage is sensitive to both N deposition and warming in N‐rich tropical soils, with interacting effects of these two global change factors. N deposition has the potential to increase total soil C stocks in tropical forests, but the long‐term stability of this added C will likely depend on future changes in temperature.     

氮沉降对热带亚热带森林土壤氮循环微生物过程的影响研究进展

近年来,高速的城市化和工业化建设导致全球大气氮沉降量逐年递增,其中热带亚热带地区氮沉降量显著高于全球平均水平,而大部分热带亚热带森林土壤趋近氮饱和状态,氮沉降增加将持续向土壤输入外源活性氮,极易导致土壤氮过剩,进而破环整个森林生态系统氮循环的平衡。我国热带亚热带地区经济发展快速,氮沉降增加导致的土壤养分失衡和林地退化等生态问题日益凸显,森林土壤氮循环对大气氮沉降的响应及适应机制已引起了学术界的广泛关注。研究表明氮循环各环节均由特定的功能微生物驱动完成,明确氮沉降增加对热带亚热带森林土壤氮循环功能微生物及其介导的关键过程的影响,对评价未来氮沉降增加背景下全球森林土壤氮循环的响应及驱动机制有重要作用,可为促进我国热带亚热带地区森林修复、生态环境的改善与提升提供科学支撑。鉴于此,本文综述了热带亚热带森林土壤氮循环主要过程（如固氮、硝化、反硝化、厌氧氨氧化等）及其功能微生物群落丰度、活性、组成等对氮沉降增加的响应,同时分析了这些功能微生物的群落特征与主要环境因子（如NH₄⁺、NO₃^-、有机碳、pH、含水量等）的关联性。在此基础上探讨了氮沉降增加下功能微生物对热带亚热带森林土壤氮循环的调控作用,重点探讨了功能微生物如何通过改变丰度与群落组成而影响氮循环过程,并对目前研究中存在的主要问题与未来研究重点进行了简要剖析。     

Carbon and nutrient limitation of soil microorganisms and microbial grazers in a tropical montane rain forest

We investigated the role of carbon, nitrogen and phosphorus as limiting factors of microorganisms and microbial grazers (testate amoebae) in a montane tropical rain forest in southern Ecuador. Carbon (as glucose), nitrogen (as NH₄NO₃) and phosphorus (as NaH₂PO₄) were added separately and in combination bimonthly to experimental plots for 20 months. By adding glucose and nutrients we expected to increase the growth of microorganisms as the major food resource of testate amoebae. The response of microorganisms to experimental treatments was determined by analysing microbial biomass (SIR), fungal biomass and microbial community composition as measured by phospholipid fatty acids (PLFAs). We hypothesized that the response of testate amoebae is closely linked to that of microorganisms. Carbon addition strongly increased ergosterol concentration and, less pronounced, the amount of linoleic acid as fungal biomarker, suggesting that saprotrophic fungi are limited by carbon. Microbial biomass and ergosterol concentrations reached a maximum in the combined treatment with C, N and P indicating that both N and P also were in short supply. In contrast to saprotrophic fungi and microorganisms in total, testate amoebae suffered from the addition of C and reached maximum density by the addition of N. The results indicate that saprotrophic fungi in tropical montane rain forests are mainly limited by carbon whereas gram positive and negative bacteria benefit from increased availability of P. Testate amoebae suffered from increased dominance of saprotrophic fungi in glucose treatments but benefited from increased supply of N. The results show that testate amoebae of tropical montane rain forests are controlled by bottom–up forces relying on specific food resources rather than the amount of bacterial biomass with saprotrophic fungi functioning as major antagonists. Compared to temperate systems microbial food webs in tropical forests therefore may be much more complex than previously assumed with trophic links being rather specific and antagonistic interactions overriding trophic interactions.     

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.     

Interactive Effects of Nitrogen and Phosphorus on Soil Microbial Communities in a Tropical Forest

Elevated nitrogen (N) deposition in humid tropical regions may exacerbate phosphorus (P) deficiency in forests on highly weathered soils. However, it is not clear how P availability affects soil microbes and soil carbon (C), or how P processes interact with N deposition in tropical forests. We examined the effects of N and P additions on soil microbes and soil C pools in a N-saturated old-growth tropical forest in southern China to test the hypotheses that (1) N and P addition will have opposing effects on soil microbial biomass and activity, (2) N and P addition will alter the composition of the microbial community, (3) the addition of N and P will have interactive effects on soil microbes and (4) addition-mediated changes in microbial communities would feed back on soil C pools. Phospholipid fatty acid (PLFA) analysis was used to quantify the soil microbial community following four treatments: Control, N addition (15 g N m⁻² yr⁻¹), P addition (15 g P m⁻² yr⁻¹), and N&P addition (15 g N m⁻² yr⁻¹ plus 15 g P m⁻² yr⁻¹). These were applied from 2007 to 2011. Whereas additions of P increased soil microbial biomass, additions of N reduced soil microbial biomass. These effects, however, were transient, disappearing over longer periods. Moreover, N additions significantly increased relative abundance of fungal PLFAs and P additions significantly increased relative abundance of arbuscular mycorrhizal (AM) fungi PLFAs. Nitrogen addition had a negative effect on light fraction C, but no effect on heavy fraction C and total soil C. In contrast, P addition significantly decreased both light fraction C and total soil C. However, there were no interactions between N addition and P addition on soil microbes. Our results suggest that these nutrients are not co-limiting, and that P rather than N is limiting in this tropical forest.     

Priming effect and microbial diversity in ecosystem functioning and response to global change: a modeling approach using the SYMPHONY model

Integration of the priming effect (PE) in ecosystem models is crucial to better predict the consequences of global change on ecosystem carbon (C) dynamics and its feedbacks on climate. Over the last decade, many attempts have been made to model PE in soil. However, PE has not yet been incorporated into any ecosystem models. Here, we build plant/soil models to explore how PE and microbial diversity influence soil/plant interactions and ecosystem C and nitrogen (N) dynamics in response to global change (elevated CO₂ and atmospheric N depositions). Our results show that plant persistence, soil organic matter (SOM) accumulation, and low N leaching in undisturbed ecosystems relies on a fine adjustment of microbial N mineralization to plant N uptake. This adjustment can be modeled in the SYMPHONY model by considering the destruction of SOM through PE, and the interactions between two microbial functional groups: SOM decomposers and SOM builders. After estimation of parameters, SYMPHONY provided realistic predictions on forage production, soil C storage and N leaching for a permanent grassland. Consistent with recent observations, SYMPHONY predicted a CO₂‐induced modification of soil microbial communities leading to an intensification of SOM mineralization and a decrease in the soil C stock. SYMPHONY also indicated that atmospheric N deposition may promote SOM accumulation via changes in the structure and metabolic activities of microbial communities. Collectively, these results suggest that the PE and functional role of microbial diversity may be incorporated in ecosystem models with a few additional parameters, improving accuracy of predictions.     

Microbial mechanisms of carbon priming effects revealed during the interaction of crop residue and nutrient inputs in contrasting soils

Agronomic practices such as crop residue return and additional nutrient supply are recommended to increase soil organic carbon (SOC) in arable farmlands. However, changes in the priming effect (PE) on native SOC mineralization in response to integrated inputs of residue and nutrients are not fully known. This knowledge gap along with a lack of understanding of microbial mechanisms hinders the ability to constrain models and to reduce the uncertainty to predict carbon (C) sequestration potential. Using a ¹³C‐labeled wheat residue, this 126‐day incubation study examined the dominant microbial mechanisms that underpin the PE response to inputs of wheat residue and nutrients (nitrogen, phosphorus and sulfur) in two contrasting soils. The residue input caused positive PE through “co‐metabolism,” supported by increased microbial biomass, C and nitrogen (N) extracellular enzyme activities (EEAs), and gene abundance of certain microbial taxa (Eubacteria, β‐Proteobacteria, Acidobacteria, and Fungi). The residue input could have induced nutrient limitation, causing an increase in the PE via “microbial nutrient mining” of native soil organic matter, as suggested by the low C‐to‐nutrient stoichiometry of EEAs. At the high residue, exogenous nutrient supply (cf. no‐nutrient) initially decreased positive PE by alleviating nutrient mining, which was supported by the low gene abundance of Eubacteria and Fungi. However, after an initial decrease in PE at the high residue with nutrients, the PE increased to the same magnitude as without nutrients over time. This suggests the dominance of “microbial stoichiometry decomposition,” supported by higher microbial biomass and EEAs, while Eubacteria and Fungi increased over time, at the high residue with nutrients cf. no‐nutrient in both soils. Our study provides novel evidence that different microbial mechanisms operate simultaneously depending on organic C and nutrient availability in a residue‐amended soil. Our results have consequences for SOC modeling and integrated nutrient management employed to increase SOC in arable farmlands.     

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.     

Nitrogen addition reduces soil respiration in a mature tropical forest in southern China

Response of soil respiration (CO₂ emission) to simulated nitrogen (N) deposition in a mature tropical forest in southern China was studied from October 2005 to September 2006. The objective was to test the hypothesis that N addition would reduce soil respiration in N saturated tropical forests. Static chamber and gas chromatography techniques were used to quantify the soil respiration, following four‐levels of N treatments (Control, no N addition; Low‐N, 5 g N m^?2 yr^?1; Medium‐N, 10 g N m^?2 yr^?1; and High‐N, 15 g N m^?2 yr^?1 experimental inputs), which had been applied for 26 months before and continued throughout the respiration measurement period. Results showed that soil respiration exhibited a strong seasonal pattern, with the highest rates found in the warm and wet growing season (April–September) and the lowest rates in the dry dormant season (December–February). Soil respiration rates showed a significant positive exponential relationship with soil temperature, whereas soil moisture only affect soil respiration at dry conditions in the dormant season. Annual accumulative soil respiration was 601±30 g CO₂‐C m^?2 yr^?1 in the Controls. Annual mean soil respiration rate in the Control, Low‐N and Medium‐N treatments (69±3, 72±3 and 63±1 mg CO₂‐C m^?2 h^?1, respectively) did not differ significantly, whereas it was 14% lower in the High‐N treatment (58±3 mg CO₂‐C m^?2 h^?1) compared with the Control treatment, also the temperature sensitivity of respiration, Q₁₀ was reduced from 2.6 in the Control with 2.2 in the High‐N treatment. The decrease in soil respiration occurred in the warm and wet growing season and were correlated with a decrease in soil microbial activities and in fine root biomass in the N‐treated plots. Our results suggest that response of soil respiration to atmospheric N deposition in tropical forests is a decline, but it may vary depending on the rate of N deposition.     

氮沉降增加情景下植物-土壤-微生物交互对自然生态系统土壤有机碳的调控研究进展

大气氮沉降增加倾向于促进受氮限制陆地生态系统地上生物量,但是对地下碳过程和土壤碳截存的影响结果迥异,导致陆地生态系统“氮促碳汇”的评估存在很大的不确定性。大气氮沉降输入直接影响微生物活性或间接影响底物质量,改变凋落物和土壤有机质(SOM)的分解速率和分解程度,进而影响土壤有机碳(SOC)的积累与损耗过程。过去相关研究主要集中在土壤碳转化过程和碳储量动态方面,缺乏植物-微生物-SOM交互作用的理解,对土壤碳截存调控的生物化学和微生物学机理尚不清楚。本文以地下碳循环过程为主线,分别综述了氮沉降增加对植物地下碳分配、SOC激发效应、微生物群落碳代谢过程的影响,深入分析SOM化学稳定性与微生物群落动态的关系。该领域研究的薄弱环节体现在:(1)增氮倾向于降低根系的生长和周转,对根际沉积碳分配(数量和格局)的影响及驱动因素不明确;(2)虽然认识到氮素有效性影响土壤激发效应的方向和强度,但是氧化态NO-3和还原态NH+4输入对有机质激发效应的差异性影响及潜在机理知之甚少;(3)微生物碳利用效率(CUE)是微生物群落碳代谢的关键表征,能够很好地解释土壤碳的积累与损耗过程;由于缺乏适宜的测定方法,难以准确量化土壤微生物的CUE及微生物生物量的周转时间;(4)增氮会抑制土壤真菌群落及其胞外酶活性,对细菌群落组成的影响尚未定论,有关SOM化学质量与土壤微生物群落活性、组成之间的耦合关系尚不清楚。未来研究应基于长期的氮添加控制实验平台,结合碳氧稳定性同位素示踪、有机质化学、分子生物学和宏基因组学等方法,深入分析植物同化碳的地下分配规律、微生物碳代谢和周转、有机质化学结构与功能微生物群落的耦合关系等关键环节。上述研究将有助于揭示植物-土壤-微生物交互作用对SOC动态的调控机制,完善陆地生态系统碳-氮耦合循环模型,有效降低区域陆地碳汇评估的不确定性,并可为陆地生态系统应对全球变化提供科学依据。     

Exotic grasses and nitrate enrichment alter soil carbon cycling along an urban–rural tropical forest gradient

Urban areas are expanding rapidly in tropical regions, with potential to alter ecosystem dynamics. In particular, exotic grasses and atmospheric nitrogen (N) deposition simultaneously affect tropical urbanized landscapes, with unknown effects on properties like soil carbon (C) storage. We hypothesized that (H1) soil nitrate (NO₃^?) is elevated nearer to the urban core, reflecting N deposition gradients. (H2) Exotic grasslands have elevated soil NO₃^? and decreased soil C relative to secondary forests, with higher N promoting decomposer activity. (H3) Exotic grasslands have greater seasonality in soil NO₃^? vs. secondary forests, due to higher sensitivity of grassland soil moisture to rainfall. We predicted that NO₃^? would be positively related to dissolved organic C (DOC) production via changes in decomposer activity. We measured six paired grassland/secondary forest sites along a tropical urban‐to‐rural gradient during the three dominant seasons (hurricane, dry, and early wet). We found that (1) soil NO₃^? was generally elevated nearer to the urban core, with particularly clear spatial trends for grasslands. (2) Exotic grasslands had lower soil C than secondary forests, which was related to elevated decomposer enzyme activities and soil respiration. Unexpectedly, soil NO₃^? was negatively related to enzyme activities, and was lower in grasslands than forests. (3) Grasslands had greater soil NO₃^? seasonality vs. forests, but this was not strongly linked to shifts in soil moisture or DOC. Our results suggest that exotic grasses in tropical regions are likely to drastically reduce soil C storage, but that N deposition may have an opposite effect via suppression of enzyme activities. However, soil NO₃^? accumulation here was higher in urban forests than grasslands, potentially related to of aboveground N interception. Net urban effects on C storage across tropical landscapes will likely vary depending on the mosaic of grass cover, rates of N deposition, and responses by local decomposer communities.     

Effects of Increased Nitrogen Availability on C and N Cycles in Tropical Forests: A Meta-Analysis

Atmospheric N deposition is predicted to increase four times over its current status in tropical forests by 2030. Our ability to understand the effects of N enrichment on C and N cycles is being challenged by the large heterogeneity of the tropical forest biome. The specific response will depend on the forest’s nutrient status; however, few studies of N addition appear to incorporate the nutrient status in tropical forests, possibly due to difficulties in explaining how this status is maintained. We used a meta-analysis to explore the consequences of the N enrichment on C and N cycles in tropical montane and lowland forests. We tracked changes in aboveground and belowground plant C and N and in mineral soil in response to N addition. We found an increasing trend of plant biomass in montane forests, but not in lowland forests, as well as a greater increase in NO emission in montane forest compared with lowland forest. The N₂O and NO emission increase in both forest; however, the N₂O increase in lowland forest was significantly even at first time N addition. The NO emission increase showed be greater at first term compared with long term N addition. Moreover, the increase in total soil N, ammonium, microbial N, and dissolved N concentration under N enrichment indicates a rich N status of lowland forests. The available evidence of N addition experiments shows that the lowland forest is richer in N than montane forests. Finally, the greater increase in N leaching and N gas emission highlights the importance of study the N deposition effect on the global climate change.     

Global forest carbon uptake due to nitrogen and phosphorus deposition from 1850 to 2100

Spatial patterns and temporal trends of nitrogen (N) and phosphorus (P) deposition are important for quantifying their impact on forest carbon (C) uptake. In a first step, we modeled historical and future change in the global distributions of the atmospheric deposition of N and P from the dry and wet deposition of aerosols and gases containing N and P. Future projections were compared between two scenarios with contrasting aerosol emissions. Modeled fields of N and P deposition and P concentration were evaluated using globally distributed in situ measurements. N deposition peaked around 1990 in European forests and around 2010 in East Asian forests, and both increased sevenfold relative to 1850. P deposition peaked around 2010 in South Asian forests and increased 3.5‐fold relative to 1850. In a second step, we estimated the change in C storage in forests due to the fertilization by deposited N and P (?C_{ν dep}), based on the retention of deposited nutrients, their allocation within plants, and C:N and C:P stoichiometry. ?C_{ν dep} for 1997–2013 was estimated to be 0.27 ± 0.13 Pg C year^?1 from N and 0.054 ± 0.10 Pg C year^?1 from P, contributing 9% and 2% of the terrestrial C sink, respectively. Sensitivity tests show that uncertainty of ?C_{ν dep} was larger from P than from N, mainly due to uncertainty in the fraction of deposited P that is fixed by soil. ?C_{P dep} was exceeded by ?C_{N dep} over 1960–2007 in a large area of East Asian and West European forests due to a faster growth in N deposition than P. Our results suggest a significant contribution of anthropogenic P deposition to C storage, and additional sources of N are needed to support C storage by P in some Asian tropical forests where the deposition rate increased even faster for P than for N.     

Soil microbial community indices as predictors of soil solution chemistry and N leaching in Picea abies (L.) Karst. forests in S. Sweden

High rates of inorganic nitrogen (N) deposition or internal N turnover increases the risks of N loss from forests with negative effects on stream water quality. We hypothesized that soil fungi may be more important N sinks than bacteria, and thus examined the impact of soil microbial community composition on N leaching from forests. We studied 19 spruce stands to examine relationships between microbial community composition, stem growth, soil-, and lysimeter-collected soil solution characteristics, and N leaching. We used nitrate concentration in the soil solution below the rooting zone as an N leaching index and phospholipid fatty acid (PLFA) analysis for characterisation of microbial communities. Microbial community composition in the organic horizon and soil solution chemistry below the rooting zone was highly correlated. Stands with low concentrations of nitrate (NO₃ ^?) and aluminium (Al) had higher fungi: bacteria ratio compared with stands with higher concentrations of NO₃ ^? and Al. Stem growth and fungi: bacteria ratio explained 70 % of the variation in N and Al leaching. We identified three microbial predictors of variation in soil solution chemistry, of which the fungi: bacteria was the strongest. The other two were putative indicators of microbial C limitation, a condition known to stimulate N mineralisation and nitrification.