Different temperature sensitivity and kinetics of soil enzymes indicate seasonal shifts in C,N and P nutrient stoichiometry in acid forest soil

Acid forest soils in the Bohemian Forest in Central Europe are biogeochemically imbalanced in organic C, N and P processing. We hypothesized that these imbalances can be due to different temperature sensitivities of soil enzyme activities and their affinities to substrate in litter and organic soil horizons. We measured potential activities of five main soil enzymes (β-glucosidase, cellobiohydrolase, Leu-aminopeptidase, Ala-aminopeptidase, and phosphatase) responsible for organic carbon, nitrogen and phosphorus acquisition. We also modeled potential in situ enzyme activities and nutrient release based on continuous in situ temperature measurements. We determined basic kinetic parameters (K_m, V_max), enzyme efficiencies (k_cat) and temperature sensitivities (E_a and Q₁₀) according to Michaelis–Menten kinetic and modified Arrhenius models. Our results showed significant differences in substrate affinities between the litter and organic soil horizons. Higher aminopeptidase affinity (lower K_m) in the litter soil horizon can lead to leaching of peptidic compounds to lower soil horizons. β-Glucosidase and phosphatase showed high temperature response following the Arrhenius model. However, both aminopeptidases showed no or even decreased activity with increasing temperature. The aminopeptidase temperature insensitivity means that peptidic compounds are degraded at the same or even lower rate in warmer and colder periods of the year in acid forest soils. This imbalance results in different release of available nutrients from plant litter and soil organic matter which may affect bacterial and fungal community composition and nutrient leaching from these ecosystems.     

A mechanism of abiotic immobilization of nitrate in forest ecosystems: the ferrous wheel hypothesis

Forest soils, rather than woody biomass, are the dominant long‐term sink for N in forest fertilization studies and, by inference, for N from atmospheric deposition. Recent evidence of significant abiotic immobilization of inorganic‐N in forest humus layers challenges a previously widely held view that microbial processes are the dominant pathways for N immobilization in soil. Understanding the plant, microbial, and abiotic mechanisms of N immobilization in forest soils has important implications for understanding current and future carbon budgets. Abiotic immobilization of nitrate is particularly perplexing because the thermodynamics of nitrate reduction in soils are not generally favorable under oxic conditions. Here we present preliminary evidence for a testable hypothesis that explains abiotic immobilization of nitrate in forest soils. Because iron (and perhaps manganese) plays a key role as a catalyst, with Fe(II) reducing nitrate and reduced forms of carbon then regenerating Fe(II), we call this ‘the ferrous wheel hypothesis’. After nitrate is reduced to nitrite, we hypothesize that nitrite reacts with dissolved organic matter through nitration and nitrosation of aromatic ring structures, thus producing dissolved organic nitrogen (DON). In addition to ignorance about mechanisms of DON production, little is known about DON dynamics in soil and its fate within ecosystems. Evidence from leaching and watershed studies suggests that DON production and consumption may be largely uncoupled from seasonal biological processes, although biological processes ultimately produce the DOC and reducing power that affect DON formation and the entire N cycle. The ferrous wheel hypothesis includes both biological and abiological processes, but the reducing power of plant‐derived organic matter may build up over seasons and years while the abiotic reduction of nitrate and reaction of organic matter with nitrite may occur in a matter of seconds after nitrate enters the soil solution.     

Anthropogenic N deposition increases soil organic matter accumulation without altering its biochemical composition

Accumulating evidence indicates that future rates of atmospheric N deposition have the potential to increase soil C storage by reducing the decay of plant litter and soil organic matter (SOM). Although the microbial mechanism underlying this response is not well understood, a decline in decay could alter the amount, as well as biochemical composition of SOM. Here, we used size‐density fractionation and solid‐state ¹³C‐NMR spectroscopy to explore the extent to which declines in microbial decay in a long‐term (ca. 20 yrs.) N deposition experiment have altered the biochemical composition of forest floor, bulk mineral soil, as well as free and occluded particulate organic matter. Significant amounts of organic matter have accumulated in occluded particulate organic matter (~20%; oPOM); however, experimental N deposition had not altered the abundance of carboxyl, aryl, alkyl, or O/N‐alkyl C in forest floor, bulk mineral soil, or any soil fraction. These observations suggest that biochemically equivalent organic matter has accumulated in oPOM at a greater rate under experimental N deposition, relative to the ambient treatment. Although we do not understand the process by which experimental N deposition has fostered the occlusion of organic matter by mineral soil particles, our results highlight the importance of interactions among the products of microbial decay and the chemical and physical properties of silt and clay particles that occlude organic matter from microbial attack. Because oPOM can reside in soils for decades to centuries, organic matter accumulating under future rates of anthropogenic N deposition could remain in soil for long periods of time. If temperate forest soils in the Northern Hemisphere respond like those in our experiment, then unabated deposition of anthropogenic N from the atmosphere has the potential to foster greater soil C storage, especially in fine‐texture forest soils.     

碳氮添加对草地土壤有机碳氮磷含量及相关酶活性的影响

草地土壤有机碳(C)、氮(N)、磷(P)等养分含量和酶活性对草地生态系统能量和养分的保持和供应具有重要作用.氮沉降对草地生态系统土壤有机养分及酶活性产生影响的结果不一致性,碳的同步添加是否会缓解氮沉降造成的负面影响仍不清楚,需要深入探讨.本研究以在内蒙古呼伦贝尔草原开展的碳(葡萄糖)、氮(尿素)添加试验样地为依托(始于2014年5月),探讨碳、氮添加对草地土壤C、N、P含量及相关酶活性的影响及其机制.试验分别设N₀(对照)、N₂₅(施氮25 kg·hm^-2·a^-1)、N₅₀(50 kg·hm^-2·a^-1)、N₁₀₀(100 kg·hm^-2·a^-1)、N₂₀₀(200 kg·hm^-2·a^-1)共5个N添加主处理,C₀(对照)、C₂₅₀(施碳250 kg·hm^-2·a^-1)、C₅₀₀(500 kg·hm^-2·a^-1)3个碳添加副处理,试验样品采于2016年8月.结果表明:高氮添加显著抑制脱氢酶(DHA)和β-1,4-N-乙酰氨基葡糖苷酶(NAG)活性,与对照相比,其活性分别降低22.3%和12.5%;而氮添加对土壤有机N含量无显著影响,使有机C和有机P含量分别减少6.6%和14.5%.高碳添加缓解了土壤微生物的碳限制,使得脱氢酶(DHA)、β-葡糖苷酶(BG)活性及土壤有机N、有机P含量分别增加15.1%、12.2%、1.9%、2.6%.研究表明,长期过量氮输入抑制土壤微生物活性,造成土壤有机C、有机P的减少,而碳添加提高了微生物及酶活性,使土壤有机N、P含量增加.碳氮耦合添加对草地土壤有机C、N、P的持续供应具有重要意义.     

Assessing the microbiological,biochemical, soil-physical and hydrological effects of amelioration of degraded soils in semiarid Spain

The way of improving degraded soils fertility and particularly of improving its microbial activity is to add “young” exogenous organic matter that contribute to provide labile organic matter to stimulate the life of the microorganisms existing in the soil. This organic matter will also improve both the retention and hydraulic characteristics of the degraded soils, all this contributing to soil restoration. In this study, the microbiological, biochemical, soil-physical and hydrological effects of the addition of a municipal solid waste compost to a degraded soil in El Campello, SE Spain were evaluated in a field experiment. Soil samples from experimental plots were analyzed 6 and 18 months after soil amendment. In both sampling time treated plots showed significantly higher microbial biomass carbon and dehydrogenase activity values than control, indicating that soil microbial population’s development and activity were stimulated by compost addition, this effect being not ephemeral but lasting in the time. Soil urease activity was not affected by compost addition while protease hydrolysing N-α-benzoil-L-argininamide (BAA) activity was strongly stimulated by the incorporation of compost into the soils. Phosphatase and β-glucosidase activities were also stimulated by the organic amendment, this stimulation being particularly noticeable 18 months after the compost addition. Nevertheless, this increase in soil microbial populations and activity did not result in an increase in soil aggregation and hydrological parameters. This can be due to the high content of carbonates and Ca²⁺ ions in these calcareous soils, that lead to an initially high content of water-stable macroaggregates. Presented at the International Conference on Bioclimatology and Natural Hazards, Poľana nad Detvou, Slovakia, 17–20 September 2007.     

Nutrient dynamics along a precipitation gradient in European beech forests

Precipitation as a key determinant of forest productivity influences forest ecosystems also indirectly through alteration of the nutrient status of the soil, but this interaction is not well understood. Along a steep precipitation gradient, we studied the consequences of reduced precipitation for the soil and biomass nutrient pools and dynamics in 14 mature European beech (Fagus sylvatica L.) forests on Triassic sandstone. We tested the hypotheses that lowered summer precipitation (1) is associated with less acid soils and (2) a reduced accumulation of organic matter on the forest floor, and (3) reduces nutrient supply from the soil and leads to decreasing foliar and root nutrient concentrations. Soil acidity, the amount of forest floor organic matter, and the associated organic matter N and P pools decreased to about a half from wet to dry sites; the C/P and N/P ratios, but not the C/N ratio, of forest floor organic matter were reduced as well. Net N mineralization and P and K pools in the mineral soil did not change with decreasing precipitation. Foliar P and K concentrations (beech sun leaves) increased while N remained constant, resulting in decreasing foliar N/P and N/K ratios. Estimated N resorption efficiency increased toward the dry sites. We conclude that a reduction in summer rainfall significantly reduces the soil C, N and P pools but does not result in decreasing foliar N and P contents in beech. However, the decreasing foliar N/P ratios towards the dry stands indicate that the importance of P limitation for tree growth declines with decreasing precipitation.     

Nitrogen deposition and leaching from two forested catchments in Southwest China--preliminary data and research needs

Increased nitrogen deposition has resulted in increased nitrogen pools and nitrogen leaching in European and North American forest soils. The development in Asia in general, and China in particular, suggests increased deposition of reduced nitrogen from changes in agricultural practices and of oxidized nitrogen from rapid growth of the transportation sector. Decreased nitrogen retention in forested areas in the future may cause increased NO3- leaching and, thus, acidification and eutrophication in surface waters. The differences in climate, ecosystems, land use, and deposition history make direct application of knowledge from studies in Europe and North America difficult. In Southwest China the potential for nitrogen mobilization from forest soils may be high because of the warm and humid climate, resulting in high decomposition rates of soil organic matter. However, there are very few data available for quantifying the suspected potential for increased nitrogen leaching in forest ecosystems. Here we present data from two forested catchments, dominated by Masson pine (Pinus massoniana), near Guiyang and Chongqing, respectively, in Southwest China. The present nitrogen deposition is moderate, estimated in the range from 10 to 40 kg N ha(-1) year(-1). The C/N ratios of the soils are generally below 15. Nitrate concentrations in soil water are rather variable in space, with highest values of several hundred microequivalents per liter. The turnover rate of nitrogen in the forest ecosystem is quite high compared to the atmospheric deposition rate. At present, nitrate runoff from the catchments is low and intermediate in Guiyang and Chongqing, respectively. More research is needed to improve our ability to predict future nitrogen leaching from subtropical Asian coniferous forests.     

Priming and microbial nutrient limitation in lowland tropical forest soils of contrasting fertility

Priming is an increase in soil organic carbon decomposition following input of labile organic carbon. In temperate soils where biological activity is limited commonly by nitrogen availability, priming is expected to occur through microbial acquisition of nitrogen from organic matter or stimulated activity of recalcitrant-carbon degrading microorganisms. However, these priming mechanisms have not yet been assessed in strongly weathered tropical forest soils where biological activity is often limited by the availability of phosphorus. We examined whether microbial nutrient limitation or community dynamics drive priming in three lowland tropical forest soils of contrasting fertility (‘low’, ‘mid’ and ‘high’) by applying C₄-sucrose (alone or in combination with nutrients; nitrogen, phosphorus and potassium) and measuring (1) the δ¹³C-signatures in respired CO₂ and in phospholipid fatty acid (PLFA) biomarkers, and (2) the activities of enzymes involved in nitrogen (N-acetyl β-glucosaminidase), phosphorus (phosphomonoesterase) and carbon (β-glucosidase, cellobiohydrolase, xylanase, phenol oxidase) acquisition from organic compounds. Priming was constrained in part by nutrient availability, because priming was greater when sucrose was added alone compared to when added with nutrients. However, the greatest priming with sucrose addition alone was detected in the medium fertility soil. Priming occurred in parallel with stimulated activity of phosphomonoesterase and phenol oxidase (but not N-acetyl β-glucosaminidase); when sucrose was added with nutrients there were lower activities of phosphomonoesterase and phenol oxidase. There was no evidence according to PLFA δ¹³C-incorporation that priming was caused by specific groups of recalcitrant-carbon degrading microorganisms. We conclude that priming occurred in the intermediate fertility soil following microbial mineralization of organic nutrients (phosphorus in particular) and suggest that priming was constrained in the high fertility soil by high nutrient availability and in the low fertility soil by the low concentration of soil organic matter amenable to priming. This first study of priming mechanisms in tropical forest soils indicates that input of labile carbon can result in priming by microbial mineralization of organic nutrients, which has important implications for understanding the fate of organic carbon in tropical forest soils.     

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

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

Monitoring the Active Conformation of Green Fluorescent Protein (GFP) and β-Glucosidase Adsorbed on Soil Particles

In order to determine the effect of various soil components on the activity of proteins, we monitored the fluorescence and the enzymatic activity of, respectively, green fluorescent protein (GFP) and β-glucosidase adsorbed on fine soil particles. We also monitored the activity of these proteins in the presence of components that are representative of soil colloids: a montmorillonite clay, goethite and organic matter extracted from soil. Upon adsorption on clay and goethite, GFP lost its fluorescence properties while β-glucosidase suffered only a partial loss of its catalytic activity. Extractable organic matter had an inactivating role on GFP while it did not cause inactivation of β-glucosidase. When GFP and β-glucosidase adsorbed on particles from natural soil samples, their behaviour was consistent with the behaviour observed for these proteins in the presence of the separate components, suggesting that the macroscopic activity of proteins adsorbed on soil particles corresponds to an average of the activities of proteins adsorbed on a mixture of surfaces. The monitoring of the proteins on soil particles with different organic matter contents has also shown that organic matter can have different effects (protecting or inactivating) on different proteins.     

亚热带米槠天然林土壤氨氧化微生物对模拟氮沉降的响应

我国亚热带地区是全球氮沉降的热点区域。氮沉降会影响氨氧化微生物的丰度和群落结构,进而改变土壤微生物驱动的养分循环。目前对新近发现的完全氨氧化菌认识不足,极大地制约了对森林土壤氨氧化微生物响应氮沉降的整体认识。本研究以福建省三明市辛口镇格氏栲自然保护区长期模拟氮沉降处理土壤为研究对象,利用实时定量PCR方法,研究氨氧化微生物(包括氨氧化细菌AOB、氨氧化古菌AOA和完全氨氧化菌comammox Nitrospira),尤其是完全氨氧化菌的amoA基因丰度。模拟氮沉降处理包括:不添加N(CK)、低氮(添加40 kg N·hm^-2·a^-1,LN)和高氮(添加80 kg N·hm^-2·a^-1,HN)。结果表明: 8年的氮添加降低了土壤pH值和有机碳含量,提高了土壤硝态氮含量。供试土壤的AOB丰度低于检测限,无法获得目的片段。高氮处理显著提高了AOA丰度,但对完全氨氧化菌clade A和clade B丰度无显著影响。两种氮添加处理均降低了完全氨氧化菌/AOA值,表明氮添加降低了完全氨氧化菌在亚热带森林土壤氨氧化微生物类群中的相对竞争力。针对完全氨氧化菌clade A和clade B的扩增都存在非特异性产物,表明针对森林土壤的高特异性和覆盖度设计引物的必要性。Clade A和clade B丰度与总氮和铵态氮含量呈显著正相关,clade B丰度还与有机碳含量呈显著正相关。总之,模拟氮沉降提高了AOA在亚热带米槠天然林土壤硝化过程中的相对重要性,这些发现可为该地区应对全球变化和氮沉降的风险评估提供理论依据。     

Retention and leaching losses of atmospherically-derived nitrogen in the aggrading coastal watershed of Waquoit Bay,MA

Extensive areas of the eastern United States are being exposed to elevated levels of nitrogen in precipitation, with levels of inorganic N in wet deposition ranging from 5 to over 20 times preindustrial, background levels. This increase in N loading to the terrestrial system, coupled with changes in land use in coastal regions in particular, has dramatically increased the level of nutrient loading from watersheds to the point that coastal waters are today among the most intensely fertilized ecosystems on earth. Studies in upland, aggrading forests have generally found that precipitation N inputs are efficiently sequestered in forest biomass and soil organic matter. However, acidic soils, sandy, porous parent substrates, and chronic inputs of salt spray common to coastal watersheds may all reduce the potential for N sequestration by the terrestrial community.We assessed the role of coastal forests in the long-term storage and retention of atmospherically-derived N in the watersheds of Waquoit Bay, MA, an increasingly eutrophic estuary on Cape Cod, by measuring precipitation inputs, storage, and lysimeter outputs below the rooting zone in a chronosequence of sites released from agriculture at different times. Calculated annual retention efficiencies were relatively low for an N-limited, aggrading forest (40–62%), and leaching losses did not vary with site age from young pine stands to mature beech forests. Nearly all nitrogen input was retained during summer months except in months with very high rainfall events. Nitrogen was released during the dormant-season in proportion to water flux through the forest floor. The composition of lysimeter output was 76% DON, 11% NO ₃ ^– , and 13% NH ₄ ^– . Total water flux and infiltration appear to be more important determinants of N retention in this sandy, coastal site than in more upland forest ecosystems; sandy systems may inherently have a low N retention efficiency.     

佛山地区存留自然林和存留人工林的土壤无机磷组分

为探讨热带亚热带森林,尤其城市及其周边地区残存森林土壤磷的有效性,对佛山地区14个残存林(7个自然林和7个人工林)的0~3 cm和3~23 cm矿质土壤的P有效性进行研究。结果表明,铁结合态无机P和还原剂可溶解无机P是土壤无机P的主要组分。在0~3 cm矿质层中,自然林土壤铝结合态无机P、Bray 1提取无机P和总无机P含量显著高于人工林;而在3~23 cm矿质土层中,自然林土壤钙结合态无机P含量显著高于人工林。其它土壤营养指标在自然林和人工林间差异不显著。相关分析结果表明,土壤有机质含量与钙结合态无机P除外的其它无机P组分含量均成显著正相关。聚类分析结果表明14个残存林土壤P有效性可分成3组,整体上人工林土壤P有效性比自然林低。这有助于认识城市化影响下城市及其周边地区残存森林土壤营养状况及加强养分管理。     

Nitrate leaching from three afforestation chronosequences on former arable land in Denmark

In regions dominated by agricultural activities, nitrogen (N) is recognized as a major pollutant in aquatic environments. In north‐western Europe, afforestation of agricultural land is part of a strategy to improve water quality. In Denmark, former arable land has been afforested during the past 40–50 years. This study evaluated the effect of afforestation of former arable land on nitrate leaching, based on three afforestation chronosequences. Precipitation, canopy throughfall and soil water were collected and soil moisture was monitored at two Danish locations, Vestskoven (nutrient‐rich, medium deposition) and Gejlvang (nutrient‐poor, high deposition). Afforestation was performed using Norway spruce [Picea abies (Karst.) L.] and common oak (Quercus robur L.) at Vestskoven and Norway spruce at Gejlvang. The results suggest that afforestation of former arable land initially leads to lower nitrate leaching than that occurring under the former agricultural land use, and largely below the standard of 50 mg NO⁻₃ L⁻¹ for groundwater to be utilized as drinking water. Nitrate concentrations became almost negligible in forest stands of 5–20 years of age. However, after canopy closure (>20 years) nitrate concentrations below the root zone and nitrate leaching tended to increase. This was attributed to increased N deposition with increasing canopy development and decreased N demand once the most N‐rich biomass compartments had been built up. Nitrate leaching started to increase at a throughfall deposition level of about 10 kg N ha⁻¹ yr⁻¹. Compared with nutrient‐poor sandy soils, nutrient‐rich clayey soils appeared more vulnerable to disturbance of the N cycle and to increased N deposition, leading to N saturation and enhanced nitrate leaching. In approximately the first 35 years after afforestation, nitrate leaching below the root zone was generally higher below oak than below Norway spruce.     

Increased inorganic nitrogen leaching from a mountain grassland ecosystem following grazing removal: a hangover of past intensive land-use?

Heathlands and grasslands occur in montane regions, naturally or due to anthropogenic land-use. These are typically nutrient-poor but exposure to elevated nitrogen deposition and intensive livestock grazing causes large-scale ecological change. We studied the long-term implications of grazing removal on soil and drainage water biogeochemistry and the implications for nitrogen cycling in 50-year replicated grazing exclosures on a montane grassland exposed to high rates of ambient nitrogen deposition. Evidence of ‘ecosystem recovery’ represented by successional change from graminoid to shrub-dominance after cessation of grazing was not reflected in the soil biogeochemistry. Cessation of grazing had a negative impact, with increased soil extractable and soil solution nitrate concentrations; an apparent shift towards a more nitrogen-rich, bacterially dominated microbial community; and the acidification of soils and leachate. The increase in nitrate leaching appears to have been counterbalanced by a decrease in dissolved organic nitrogen leaching, approximately maintaining the overall nitrogen balance of the system, whilst apparently altering ecosystem functioning. High rates of organic matter cycling and inorganic nitrogen uptake in grazed grassland may have sustained ecosystem N limitation under elevated nitrogen deposition. Grazing removal caused long-term over-supply of nitrogen from mineralisation of enriched organic matter, exacerbated by continued high nitrogen deposition, exceeding the uptake demand of heath vegetation and resulting in nitrification and nitrate leaching. This disequilibrium between vegetation and soil following grazing removal has implications for restoration after periods of intensive grazing. Grazing may not simply leave a legacy of nutrient enrichment but its cessation may trigger nitrogen saturation and soil and freshwater eutrophication and acidification which counteract the immediate benefits of natural vegetation recovery. Long term, nitrogen saturation of abandoned grasslands is likely to reduce ecosystem resilience to invasion by nitrophilous species, pathogen attack and vulnerability to environmental pressures such as climate change. We conclude that partial and/or phased reduction in grazing levels may permit the more synchronised recovery of soils and vegetation, thereby avoiding imbalances between nitrogen supply and nitrogen demand and detrimental ecological effects.     

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

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

Soil organic matter and litter chemistry response to experimental N deposition in northern temperate deciduous forest ecosystems

The effects of atmospheric nitrogen (N) deposition on organic matter decomposition vary with the biochemical characteristics of plant litter. At the ecosystem‐scale, net effects are difficult to predict because various soil organic matter (SOM) fractions may respond differentially. We investigated the relationship between SOM chemistry and microbial activity in three northern deciduous forest ecosystems that have been subjected to experimental N addition for 2 years. Extractable dissolved organic carbon (DOC), DOC aromaticity, C : N ratio, and functional group distribution, measured by Fourier transform infrared spectra (FTIR), were analyzed for litter and SOM. The largest biochemical changes were found in the sugar maple–basswood (SMBW) and black oak–white oak (BOWO) ecosystems. SMBW litter from the N addition treatment had less aromaticity, higher C : N ratios, and lower saturated carbon, lower carbonyl carbon, and higher carboxylates than controls; BOWO litter showed opposite trends, except for carbonyl and carboxylate contents. Litter from the sugar maple–red oak (SMRO) ecosystem had a lower C : N ratio, but no change in DOC aromaticity. For SOM, the C : N ratio increased with N addition in SMBW and SMRO ecosystems, but decreased in BOWO; N addition did not affect the aromaticity of DOC extracted from mineral soil. All ecosystems showed increases in extractable DOC from both litter and soil in response to N treatment. The biochemical changes are consistent with the divergent microbial responses observed in these systems. Extracellular oxidative enzyme activity has declined in the BOWO and SMRO ecosystems while activity in the SMBW ecosystem, particularly in the litter horizon, has increased. In all systems, enzyme activities associated with the hydrolysis and oxidation of polysaccharides have increased. At the ecosystem scale, the biochemical characteristics of the dominant litter appear to modulate the effects of N deposition on organic matter dynamics.     

Large Loss of Dissolved Organic Nitrogen from Nitrogen-Saturated Forests in Subtropical China

Dissolved organic nitrogen (DON) has recently been recognized as an important component of terrestrial N cycling, especially under N-limited conditions; however, the effect of increased atmospheric N deposition on DON production and loss from forest soils remains controversial. Here we report DON and dissolved organic carbon (DOC) losses from forest soils receiving very high long-term ambient atmospheric N deposition with or without additional experimental N inputs, to investigate DON biogeochemistry under N-saturated conditions. We studied an old-growth forest, a young pine forest, and a young mixed pine/broadleaf forest in subtropical southern China. All three forests have previously been shown to have high nitrate (NO₃⁻) leaching losses, with the highest loss found in the old-growth forest. We hypothesized that DON leaching loss would be forest specific and that the strongest response to experimental N input would be in the N-saturated old-growth forest. Our results showed that under ambient deposition (35–50 kg N ha⁻¹ y⁻¹ as throughfall input), DON leaching below the major rooting zone in all three forests was high (6.5–16.9 kg N ha⁻¹ y⁻¹). DON leaching increased 35–162% following 2.5 years of experimental input of 50–150 kg N ha⁻¹ y⁻¹. The fertilizer-driven increase of DON leaching comprised 4–17% of the added N. A concurrent increase in DOC loss was observed only in the pine forest, even though DOC:DON ratios declined in all three forests. Our data showed that DON accounted for 23–38% of total dissolved N in leaching, highlighting that DON could be a significant pathway of N loss from forests moving toward N saturation. The most pronounced N treatment effect on DON fluxes was not found in the old-growth forest that had the highest DON loss under ambient conditions. DON leaching was highly correlated with NO₃⁻ leaching in all three forests. We hypothesize that abiotic incorporation of excess NO₃⁻ (through chemically reactive NO₂⁻) into soil organic matter and the consequent production of N-enriched dissolved organic matter is a major mechanism for the consistent and large DON loss in the N-saturated subtropical forests of southern China. Dr. YT Fang performed research, analyzed data, and wrote the paper; Prof. WX Zhu participated in the initial experimental design, analyzed data, and took part in writing the paper; Prof. P Gundersen conceived the study and took part in writing; Prof. JM Mo and Prof. GY Zhou conceived study; Prof. M Yoh analyzed part of the data and contributed to the development of DON model.     

Effects of long-term nitrogen deposition on phosphorus leaching dynamics in a mature tropical forest

Elevated anthropogenic nitrogen (N) deposition is suggested to affect ecosystem phosphorus (P) cycling through altered biotic P demand and soil acidification. To date, however, there has been little information on how long-term N deposition regulates P fluxes in tropical forests, where P is often depleted. To address this question, we conducted a long-term N addition experiment in a mature tropical forest in southern China, using the following N treatments: 0, 50, 100, and 150 kg N ha^?1 year^?1. We hypothesized that (i) tropical forest ecosystems have conservative P cycling with low P output, and (ii) long-term N addition decreases total dissolved phosphorus (TDP) leaching losses due to reduced litter decomposition rates and stimulated P sorption deriving from accelerated soil acidification. As hypothesized, we demonstrated a closed P cycling with low leaching outputs in our forest. Under experimental N addition, TDP flux in throughfall was significantly reduced, suggesting that N addition may result in a less internal P recycling. Contrary to our hypothesis, N addition did not decrease TDP leaching, despite reduced litter decomposition and accelerated soil acidification. We find that N addition might have negative impacts on biological P uptake without affecting TDP leaching, and that the amount of TDP leaching from soil could be lower than a minimum concentration for TDP retention. Overall, we conclude that long-term N deposition does not necessarily decrease P effluxes from tropical forest ecosystems with conservative P cycling.     

Impact of Agricultural Land-use Change on Carbon Storage in Boreal Alaska

Climate warming is most pronounced at high latitudes, which could result in the intensification of the extensively cultivated areas in the boreal zone and could further enhance rates of forest clearing in the coming decades. Using paired forest‐field sampling and a chronosequence approach, we investigated the effect of conversion of boreal forest to agriculture on carbon (C) and nitrogen (N) dynamics in interior Alaska. Chronosequences showed large soil C losses during the first two decades following deforestation, with mean C stocks in agricultural soils being 44% or 8.3 kg m^?2 lower than C stocks in original forest soils. This suggests that soil C losses from land‐use change in the boreal region may be greater than those in other biomes. Analyses of changes in stable C isotopes and in quality of soil organic matter showed that organic C was lost from soils by combustion of cleared forest material, decomposition of organic matter and possibly erosion. Chronosequences indicated an increase in C storage during later decades after forest clearing, with 60‐year‐old grassland showing net ecosystem C gain of 2.1 kg m^?2 over the original forest. This increase in C stock resulted probably from a combination of large C inputs from belowground biomass and low C losses due to a small original forest soil C stock and low tillage frequency. Reductions in soil N stocks caused by land‐use change were smaller than reductions in C stocks (34% or 0.31 kg m^?2), resulting in lower C/N ratios in field compared with forest mineral soils, despite the occasional incorporation of high‐C forest‐floor material into field soils. Carbon mineralization per unit of mineralized N was considerably higher in forests than in fields, which could indicate that decomposition rates are more sensitive in forest soils than in field soils to inorganic N addition (e.g. by increased N deposition from the atmosphere). If forest conversion to agriculture becomes more widespread in the boreal region, the resulting C losses (51% or 11.2 kg m^?2 at the ecosystem level in this study) will induce a positive feedback to climatic warming and additional land‐use change. However, by selecting relatively C‐poor soils and by implementing management practices that preserve C, losses of C from soils can be reduced.