The effects of warming and nitrogen addition on soil nitrogen cycling in a temperate grassland, northeastern China

Background

Both climate warming and atmospheric nitrogen (N) deposition are predicted to affect soil N cycling in terrestrial biomes over the next century. However, the interactive effects of warming and N deposition on soil N mineralization in temperate grasslands are poorly understood.

Methodology/Principal Findings

A field manipulation experiment was conducted to examine the effects of warming and N addition on soil N cycling in a temperate grassland of northeastern China from 2007 to 2009. Soil samples were incubated at a constant temperature and moisture, from samples collected in the field. The results showed that both warming and N addition significantly stimulated soil net N mineralization rate and net nitrification rate. Combined warming and N addition caused an interactive effect on N mineralization, which could be explained by the relative shift of soil microbial community structure because of fungal biomass increase and strong plant uptake of added N due to warming. Irrespective of strong intra- and inter-annual variations in soil N mineralization, the responses of N mineralization to warming and N addition did not change during the three growing seasons, suggesting independence of warming and N responses of N mineralization from precipitation variations in the temperate grassland.

Conclusions/Significance

Interactions between climate warming and N deposition on soil N cycling were significant. These findings will improve our understanding on the response of soil N cycling to the simultaneous climate change drivers in temperate grassland ecosystem.     

Effects of long-term experimental warming on plants and soil microbes in a cool temperate semi-natural grassland in Japan

To clarify the effects of long-term warming on ecosystem matter cycling, we conducted an in situ 7-year experimental warming (2009–2015) using infrared heaters in a cool temperate semi-natural grassland in Japan. We measured plant aboveground biomass, soil total C and N, soil inorganic N (NH₄ ⁺-N and NO₃ ^?-N), and soil microbial biomass for 7 years (2009–2015). We also measured heterotrophic respiration for 2 years (2013–2014) and assessed net N mineralization and nitrification in 2015. We found that warming immediately increased plant aboveground biomass, but this effect ceased in 2013. However, the soil microbial biomass was continuously depressed by warming. Soil inorganic N concentrations in warmed plots substantially increased in the later years of the experiment (2013–2015) and the potential net N mineralization rate was also higher than in the earlier years. In contrast, heterotrophic respiration decreased with warming in 2013–2014. Our observations indicate that long-term warming has a contrasting effect on plants and soil microbes. In addition, the warming could have different effects on subterranean C and N cycling. To enhance the accuracy of estimation of future climate change, it is essential to continuously observe the warming effects on ecosystems and to focus on the change in subterranean C and N cycling.     

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.     

模拟大气氮沉降对中国森林生态系统影响的研究进展

人类活动加剧了活性氮的生产和排放,并导致氮沉降日益增加并全球化。目前,人类活动对全球氮循环的干扰已经超出了地球系统安全运行的界限。中国已成为全球氮沉降的高发区域,高氮沉降已经威胁到生态系统的健康和安全,并成为生态文明建设过程中亟待理清和解决的热点问题。对国际上和中国森林生态系统模拟氮沉降研究的概况进行了综述,并从生物学和非生物学两大过程重点阐述模拟氮沉降增加对中国主要森林生态系统影响的研究进展。中国自2000年以后才开始重视大气氮沉降产生的生态环境问题,中国科学院华南植物园在国内森林生态系统模拟氮沉降试验研究上做出了开创性的贡献。模拟氮沉降研究表明,持续高氮输入将会显著改变森林生态系统的结构和功能,并威胁生态系统的健康发展,特别是处于氮沉降热点区域的中国中南部。森林生态系统的氮沉降效应依赖于系统的氮状态、土地利用历史、气候特征、林型和林龄等。最后,对未来的研究提出了一些建议,包括加强长期跟踪研究和不同气候带站点之间的联网研究,特别是在森林生态系统对长期氮沉降响应与适应的过程机制、地下碳氮吸存潜力研究、以及与其他全球变化因子的耦合研究等方面,以期为森林生态系统的可持续发展提供理论基础和管理依据。     

土壤微生物对气候变暖和大气N沉降的响应

气候变暖和大气N沉降是近一、二十年来人们非常关注的全球变化现象,它们所带来的一系列生态问题已成为全球变化研究的重要议题。它们不仅影响地上植被生长和群落组成,还直接或间接地影响土壤微生物过程,而土壤微生物对此做出的响应正是生态系统反馈过程中非常重要的环节。该文分别从气候变化对土壤微生物的影响(土壤微生物量、微生物活动和微生物群落结构)和土壤微生物对气候变化的响应(凋落物分解、养分利用与循环以及养分的固持与流失)两个角度,综述近期土壤微生物对气候变暖和大气N沉降响应与适应的研究进展。气候变暖和大气N沉降对土壤微生物的影响更多地反映在微生物群落的结构和功能上,而土壤微生物量、微生物活动和群落结构的变化又会通过改变凋落物分解、养分利用和C、N循环等重要的土壤生态系统功能和过程做出响应,形成正向或负向反馈,加强或削弱气候变化给整个陆地生态系统带来的影响。然而,到目前为止土壤微生物的响应对陆地生态系统产生的最终结果仍是未决的关键性问题。     

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。     

Elevated temperature shifts soil N cycling from microbial immobilization to enhanced mineralization,nitrification and denitrification across global terrestrial ecosystems

We assessed the response of soil microbial nitrogen (N) cycling and associated functional genes to elevated temperature at the global scale. A meta‐analysis of 1,270 observations from 134 publications indicated that elevated temperature decreased soil microbial biomass N and increased N mineralization rates, both in the presence and absence of plants. These findings infer that elevated temperature drives microbially mediated N cycling processes from dominance by anabolic to catabolic reaction processes. Elevated temperature increased soil nitrification and denitrification rates, leading to an increase in N₂O emissions of up to 227%, whether plants were present or not. Rates of N mineralization, denitrification and N₂O emission demonstrated significant positive relationships with rates of CO₂ emissions under elevated temperatures, suggesting that microbial N cycling processes were associated with enhanced microbial carbon (C) metabolism due to soil warming. The response in the abundance of relevant genes to elevated temperature was not always consistent with changes in N cycling processes. While elevated temperature increased the abundances of the nirS gene with plants and nosZ genes without plants, there was no effect on the abundances of the ammonia‐oxidizing archaea amoA gene, ammonia‐oxidizing bacteria amoA and nirK genes. This study provides the first global‐scale assessment demonstrating that elevated temperature shifts N cycling from microbial immobilization to enhanced mineralization, nitrification and denitrification in terrestrial ecosystems. These findings infer that elevated temperatures have a profound impact on global N cycling processes with implications of a positive feedback to global climate and emphasize the close linkage between soil microbial C and N cycling.     

Soil–plant N processes in a High Arctic ecosystem,NW Greenland are altered by long‐term experimental warming and higher rainfall

Rapid temperature and precipitation changes in High Arctic tundra ecosystems are altering the biogeochemical cycles of carbon (C) and nitrogen (N), but in ways that are difficult to predict. The challenge grows from the uncertainty of N cycle responses and the extent to which shifts in soil N are coupled with the C cycle and productivity of tundra systems. We used a long‐term (since 2003) experiment of summer warming and supplemental summer water additions to a High Arctic ecosystem in NW Greenland, and applied a combination of discrete sampling and in situ soil core incubations to measure C and N pools and seasonal microbial processes that might control plant‐available N. We hypothesized that elevated temperature and increased precipitation would stimulate microbial activity and net inorganic N mineralization, thereby increasing plant N‐availability through the growing season. While we did find increased N mineralization rates under both global change scenarios, water addition also significantly increased net nitrification rates, loss of NO₃^?‐N via leaching, and lowered rates of labile organic N production. We also expected the chronic warming and watering would lead to long‐term changes in soil N‐cycling that would be reflected in soil δ¹⁵N values. We found that soil δ¹⁵N decreased under the different climate change scenarios. Our results suggest that temperature accelerates biological processes and existing C and N transformations, but moisture increases soil hydraulic connectivity and so alters the pathways, and changes the fate of the products of C and N transformations. In addition, our findings indicate that warmer, wetter High Arctic tundra will be cycling N and C in ways that may transform these landscapes in part leading to greater C sequestration, but simultaneously, N losses from the upper soil profile that may be transported to depth dissolved in water and or transported off site in lateral flow.     

Warming and drought reduce temperature sensitivity of nitrogen transformations

Shifts in nitrogen (N) mineralization and nitrification rates due to global changes can influence nutrient availability, which can affect terrestrial productivity and climate change feedbacks. While many single‐factor studies have examined the effects of environmental changes on N mineralization and nitrification, few have examined these effects in a multifactor context or recorded how these effects vary seasonally. In an old‐field ecosystem in Massachusetts, USA, we investigated the combined effects of four levels of warming (up to 4 °C) and three levels of precipitation (drought, ambient, and wet) on net N mineralization, net nitrification, and potential nitrification. We also examined the treatment effects on the temperature sensitivity of net N mineralization and net nitrification and on the ratio of C mineralization to net N mineralization. During winter, freeze–thaw events, snow depth, and soil freezing depth explained little of the variation in net nitrification and N mineralization rates among treatments. During two years of treatments, warming and altered precipitation rarely influenced the rates of N cycling, and there was no evidence of a seasonal pattern in the responses. In contrast, warming and drought dramatically decreased the apparent Q₁₀ of net N mineralization and net nitrification, and the warming‐induced decrease in apparent Q₁₀ was more pronounced in ambient and wet treatments than the drought treatment. The ratio of C mineralization to net N mineralization varied over time and was sensitive to the interactive effects of warming and altered precipitation. Although many studies have found that warming tends to accelerate N cycling, our results suggest that warming can have little to no effect on N cycling in some ecosystems. Thus, ecosystem models that assume that warming will consistently increase N mineralization rates and inputs of plant‐available N may overestimate the increase in terrestrial productivity and the magnitude of an important negative feedback to climate change.     

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。     

Effects of long-term nitrogen addition on phosphorus cycling in organic soil horizons of temperate forests

High atmospheric nitrogen (N) deposition is expected to impair phosphorus (P) nutrition of temperate forest ecosystems. We examined N and P cycling in organic soil horizons of temperate forests exposed to long-term N addition in the northeastern USA and Scandinavia. We determined N and P concentrations, enzyme activities and net N and P mineralization rates in organic soil horizons of two deciduous (Harvard Forest, Bear Brook) and two coniferous (Klosterhede, Gårdsjön) forests which had received experimental inorganic N addition between 25 and 150 kg N ha^?1 year^?1 for more than 25 years. Long-term N addition increased the activity of phosphatase (+?180%) and the activity of carbon (C)- and N-acquiring enzymes (cellobiohydrolase: +?70%, chitinase: +?25%). Soil N enrichment increased the N:P ratio of organic soil horizons by up to 150%. In coniferous organic soil horizons, net N and P mineralization were small and unaffected by N addition. In deciduous organic soil horizons, net N and P mineralization rates were significantly higher than at the coniferous sites, and N addition increased net N mineralization by up to 290%. High phosphatase activities concomitant with a 40% decline in P stocks of deciduous organic soil horizons indicate increased plant P demand. In summary, projected future global increases in atmospheric N deposition may induce P limitation in deciduous forests, impairing temperate forest growth.     

Plant nutrient mobilization in temperate heathland responds to elevated CO2, temperature and drought

Temperate terrestrial ecosystems are currently exposed to increased atmospheric CO₂ and progressive climatic changes with increased temperature and periodical drought. We here present results from a field experiment, where the effects of these three main climate change related factors are investigated solely and in all combinations at a temperate heathland. Significant responses were found in the top soils below the two dominant species (Calluna vulgaris and Deschampsia flexuosa). During winter incubation, microbial immobilization of N and ammonification rate decreased in response to warming in Deschampsia soil, and microbial immobilization of N and P decreased in warmed Calluna soil. Warming tended to increase microbial N and P in Calluna but not in Deschampsia soil in fall, and more microbial C was accumulated under drought in Calluna soil. The effects of warming were often counteracted or erased when combined with CO₂ and drought. Below Deschampsia, the net nitrification rate decreased in response to drought and, while phosphorus availability and microbial P immobilization decreased, but nitrification increased in response to elevated CO₂. Furthermore, leaf litter decomposition of both species decreased in response to drought. These complex changes in availability and release of nutrients from soil organic matter turnover and mineralization in response to elevated CO₂ and climate change may influence the future plant carbon sequestration and species composition at temperate heathlands.     

Decadal-scale Dynamics of Water, Carbon and Nitrogen in a California Chaparral Ecosystem: DAYCENT Modeling Results

The Mediterranean climate, with its characteristic of dry summers and wet winters, influences the hydrologic and microbial processes that control carbon (C) and nitrogen (N) biogeochemical processes in chaparral ecosystems. These biogeochemical processes in turn determine N cycling under chronic N deposition. In order to examine connections between climate and N dynamics, we quantified decadal-scale water, C and N states and fluxes at annual, monthly and daily time steps for a California chaparral ecosystem in the Sierra Nevada using the DAYCENT model. The daily output simulations of net mineralization, stream flow and stream nitrate (NO₃⁻) export were developed for DAYCENT in order to simulate the N dynamics most appropriate for the abrupt rewetting events characteristic of Mediterranean chaparral ecosystems. Overall, the magnitude of annual modeled net N mineralization, soil and plant biomass C and N, nitrate export and gaseous N emission agreed with those of observations. Gaseous N emission was a major N loss pathway in chaparral ecosystems, in which nitric oxide (NO) is the dominant species. The modeled C and N fluxes of net primary production (NPP), N uptake and N mineralization, NO₃⁻ export and gaseous N emission showed both high inter-annual and intra-annual variability. Our simulations also showed dramatic fire effects on NPP, N uptake, N mineralization and gaseous N emission for three years of postfire. The decease in simulated soil organic C and N storages was not dramatic, but lasted a longer time. For the seasonal pattern, the predicted C and N fluxes were greatest during December to March, and lowest in the summer. The model predictions suggested that an increase in the N deposition rate would increase N losses through gaseous N emission and stream N export in the chaparral ecosystems of the Sierra Nevada due to changes in N saturation status. The model predictions could not capture stream NO₃⁻ export during most rewetting events suggesting that a dry-rewetting mechanism representing the increase in N mineralization following soil wetting needs to be incorporated into biogeochemical models of semi-arid ecosystems.     

Nitrogen deposition and forest nitrogen cycling along an urban–rural transect in southern China

There is increasing concern over the impact of atmospheric nitrogen (N) deposition on forest ecosystems in the tropical and subtropical areas. In this study, we quantified atmospheric N deposition and revealed current plant and soil N status in 14 forests along a 150 km urban to rural transect in southern China, with an emphasis on examining whether foliar δ¹⁵N can be used as an indicator of N saturation. Bulk deposition ranged from 16.2 to 38.2 kg N ha^?1 yr^?1, while the throughfall covered a larger range of 11.7–65.1 kg N ha^?1 yr^?1. Foliar N concentration, NO₃^? leaching to stream, and soil NO₃^? concentration were low and NO₃^? production was negligible in some rural forests, indicating that primary production in these forests may be limited by N supply. But all these N variables were enhanced in suburban and urban forests. Across the study transect, throughfall N input was correlated positively with soil nitrification and NO₃^? leaching to stream, and negatively with pH values in soil and stream water. Foliar δ¹⁵N was between ?6.6‰ and 0.7‰, and was negatively correlated with soil NO₃^? concentration and NO₃^? leaching to stream across the entire transect, demonstrating that an increased N supply does not necessarily increase forest δ¹⁵N values. We proposed several potential mechanism that could contribute to the δ¹⁵N pattern, including (1) increased plant uptake of ¹⁵N‐depleted soil NO₃^?, (2) foliage uptake of ¹⁵N‐depleted NH₄⁺, (3) increased utilization of soil inorganic N relative to dissolved organic N, and (4) increased fractionation during plant N uptake under higher soil N availability.     

Reduced N cycling in response to elevated CO2, warming,and drought in a Danish heathland: Synthesizing results of the CLIMAITE project after two years of treatments

Field‐scale experiments simulating realistic future climate scenarios are important tools for investigating the effects of current and future climate changes on ecosystem functioning and biogeochemical cycling. We exposed a seminatural Danish heathland ecosystem to elevated atmospheric carbon dioxide (CO₂), warming, and extended summer drought in all combinations. Here, we report on the short‐term responses of the nitrogen (N) cycle after 2 years of treatments. Elevated CO₂ significantly affected aboveground stoichiometry by increasing the carbon to nitrogen (C/N) ratios in the leaves of both co‐dominant species (Calluna vulgaris and Deschampsia flexuosa), as well as the C/N ratios of Calluna flowers and by reducing the N concentration of Deschampsia litter. Belowground, elevated CO₂ had only minor effects, whereas warming increased N turnover, as indicated by increased rates of microbial NH₄⁺ consumption, gross mineralization, potential nitrification, denitrification and N₂O emissions. Drought reduced belowground gross N mineralization and decreased fauna N mass and fauna N mineralization. Leaching was unaffected by treatments but was significantly higher across all treatments in the second year than in the much drier first year indicating that ecosystem N loss is highly sensitive to changes and variability in amount and timing of precipitation. Interactions between treatments were common and although some synergistic effects were observed, antagonism dominated the interactive responses in treatment combinations, i.e. responses were smaller in combinations than in single treatments. Nonetheless, increased C/N ratios of photosynthetic tissue in response to elevated CO₂, as well as drought‐induced decreases in litter N production and fauna N mineralization prevailed in the full treatment combination. Overall, the simulated future climate scenario therefore lead to reduced N turnover, which could act to reduce the potential growth response of plants to elevated atmospheric CO₂ concentration.     

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.     

Warming prevents the elevated CO2-induced reduction in available soil nitrogen in a temperate, perennial grassland

Rising atmospheric carbon dioxide concentration ([CO₂]) has the potential to stimulate ecosystem productivity and sink strength, reducing the effects of carbon (C) emissions on climate. In terrestrial ecosystems, increasing [CO₂] can reduce soil nitrogen (N) availability to plants, preventing the stimulation of ecosystem C assimilation; a process known as progressive N limitation. Using ion exchange membranes to assess the availability of dissolved organic N, ammonium and nitrate, we found that CO₂ enrichment in an Australian, temperate, perennial grassland did not increase plant productivity, but did reduce soil N availability, mostly by reducing nitrate availability. Importantly, the addition of 2 °C warming prevented this effect while warming without CO₂ enrichment did not significantly affect N availability. These findings indicate that warming could play an important role in the impact of [CO₂] on ecosystem N cycling, potentially overturning CO₂‐induced effects in some ecosystems.     

Resistance of soil protein depolymerization rates to eight years of elevated CO<Subscript>2</Subscript>, warming,and summer drought in a temperate heathland

Soil N availability for plants and microorganisms depends on the breakdown of soil polymers such as proteins into smaller, assimilable units by microbial extracellular enzymes. Changing climatic conditions are expected to alter protein depolymerization rates over the next decades, and thereby affect the potential for plant productivity. We here tested the effect of increased CO₂ concentration, temperature, and drought frequency on gross rates of protein depolymerization, N mineralization, microbial amino acid and ammonium uptake using ¹⁵N pool dilution assays. Soils were sampled in fall 2013 from the multifactorial climate change experiment CLIMAITE that simulates increased CO₂ concentration, temperature, and drought frequency in a fully factorial design in a temperate heathland. Eight years after treatment initiation, we found no significant effect of any climate manipulation treatment, alone or in combination, on protein depolymerization rates. Nitrogen mineralization, amino acid and ammonium uptake showed no significant individual treatment effects, but significant interactive effects of warming and drought. Combined effects of all three treatments were not significant for any of the measured parameters. Our findings therefore do not suggest an accelerated release of amino acids from soil proteins in a future climate at this site that could sustain higher plant productivity.     

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.     

Original Articles: Nitrogen Mineralization in Two Unpolluted Old-Growth Forests of Contrasting Biodiversity and Dynamics

Studies in unpolluted, old-growth forests in the coastal range of southern Chile (42°30′S) can provide a baseline for understanding how forest ecosystems are changing due to the acceleration of nitrogen (N) inputs that has taken place over the last century. Chilean temperate forests, in contrast to their northern hemisphere counterparts, exhibit extremely low losses of inorganic N to stream waters. The objectives of this study were (a) to determine whether low inorganic N outputs in these forests were due to low rates of N mineralization or nitrification, and (b) to examine how biodiversity (defined as number of dominant tree species) and forest structure influence N mineralization and overall patterns of N cycling. Studies were conducted in a species-poor, conifer-dominated (Fitzroya cupressoides) forest with an even-aged canopy, and in a mixed-angiosperm (Nothofagus nitida) forest with a floristically more diverse and unstable canopy. Nitrogen mineralization rates measured in laboratory assays varied seasonally, reaching 6.0 μg N/g DW/day in both forests during late summer. Higher values were related to higher microbial activity, larger pools of labile inorganic N, and increased fine litter inputs. Field assays, conducted monthly, indicated positive net flux from N mineralization mainly from December to January in both forests. Annual net flux of N from mineralization varied from 20 to 23 kg/ha/year for the Fitzroya forest and from 31 to 37 kg/ha/year for the Nothofagus forest. Despite low losses of inorganic N to streams, N mineralization and nitrification are not inhibited in these forests, implying the existence of strong sinks for NO₃ ⁻ in the ecosystem. Field N mineralization rates were two times higher in the Nothofagus forest than in the Fitzroya forest, and correlated with greater N input via litterfall, slightly higher soil pH, and narrower carbon (C)–nitrogen ratios of soils and litter in the former. Differences in N mineralization between the two forest types are attributed to differences in biotic structure, stand dynamics, and site factors. Median values of net N mineralization rates in these southern hemisphere forests were lower than median rates for forests in industrialized regions of North America, such as the eastern and central USA. We suggest that these high N mineralization rates may be a consequence of enhanced atmospheric N deposition.