Does Atmospheric NO<Stack><Subscript>3</Subscript><Superscript>−</Superscript></Stack> Deposition Alter the Abundance and Activity of Ligninolytic Fungi in Forest Soils?

Although field studies have demonstrated an ecosystem-specific effect of experimental atmospheric nitrogen (N) deposition on litter decomposition, a mechanistic understanding of how ligninolytic microbial communities respond to atmospheric deposition is lacking. Because high levels of inorganic N suppress lignin decomposition by some basidiomycetes, it is plausible that the abundance and activity of these key microorganisms underlies differential ecosystem responses of decomposition to atmospheric N deposition. We hypothesize that: (a) atmospheric N deposition will cause an ecosystem-specific reduction in basidiomycete activity and abundance with greatest decreases in ecosystems with lignin-rich forest litter and (b) the abundance of lignin degrading basidiomycetes will be positively correlated with ligninolytic enzyme activity. To test these hypotheses, we measured the effects of experimental N deposition on the potential activity of phenol oxidase enzymes, and the abundance of basidiomycete genes encoding laccase, a primary phenol oxidase enzyme, in three hardwood forests spanning a range of leaf litter lignin content. The black oak-white oak (BOWO) contains high lignin litter, the sugar maple-basswood (SMBW) has low lignin litter, and the sugar maple-red oak (SMRO) is intermediate. An ecosystem by N deposition interaction significantly influenced phenol oxidase activity in the surface soil (P = 0.05), where phenol oxidase activity decreased with increasing experimental N deposition in the BOWO ecosystem. No consistent response to N deposition was evident for surface soil phenol oxidase activity within either the SMRO or SMBW ecosystem. This interaction did not influence laccase gene abundance. Instead, basidiomycete laccase gene abundance was reduced by experimental N deposition (main effect) in surface soil. There was only a weak correlation between basidiomycete laccase gene abundance and potential phenol oxidase enzyme activity, suggesting that the abundance of organisms possessing laccase genes may not control phenol oxidase activity in soil. Our results suggest that the regulation of laccase gene expression may mediate the decomposition response to atmospheric N deposition.     

氮、磷养分有效性对森林凋落物分解的影响研究进展

通过对相关研究文献的综述结果表明,氮(N)和磷(P)是构成蛋白质和遗传物质的两种重要组成元素,限制森林生产力和其他生态系统过程,对凋落物分解产生深刻影响。大量的凋落物分解试验发现在土壤N有效性较低的温带和北方森林,凋落物分解速率常与底物初始N浓度、木质素/N比等有很好的相关关系,也受外源N输入的影响;而在土壤高度风化的热带亚热带森林生态系统中,P可能是比N更为重要的分解限制因子。然而控制试验表明,N、P添加对凋落物分解速率的影响并不一致,既有促进效应也有抑制效应。为了深入揭示N、P养分有效性对凋落物分解的调控机制,"底物的C、N化学计量学"假说、"微生物的N开采"假说以及养分平衡的理论都常被用于解释凋落物分解速率的变化。由于微生物分解者具有较为稳定的C、N、P等养分需求比例,在不同的养分供应的周围环境中会体现出不同的活性,某种最缺乏的养分可能就是分解的最重要限制因子。未来的凋落物分解研究,应延长实验时间、加强室内和野外不同条件下的N、P等养分添加控制试验,探讨驱动分解进程的微生物群落结构和酶活性的变化。     

Interactions between Litter Lignin and Nitrogenitter Lignin and Soil Nitrogen Availability during Leaf Litter Decomposition in a Hawaiian Montane Forest

Previous work in a young Hawaiian forest has shown that nitrogen (N) limits aboveground net primary production (ANPP) more strongly than it does decomposition, despite low soil N availability. In this study, I determined whether (a) poor litter C quality (that is, high litter lignin) poses an overriding constraint on decomposition, preventing decomposers from responding to added N, or (b) high N levels inhibit lignin degradation, lessening the effects of added N on decomposition overall. I obtained leaf litter from one species, Metrosideros polymorpha, which dominates a range of sites in the Hawaiian Islands and whose litter lignin concentration declines with decreasing precipitation. Litter from three dry sites had lignin concentrations of 12% or less, whereas litter from two wet sites, including the study site, had lignin concentrations of more than 18%. This litter was deployed 2.5 years in a common site in control plots (receiving no added nutrients) and in N-fertilized plots. Nitrogen fertilization stimulated decomposition of the low-lignin litter types more than that of the high-lignin litter types. However, in contrast to results from temperate forests, N did not inhibit lignin decomposition. Rather, lignin decay increased with added N, suggesting that the small effect of N on decomposition at this site results from limitation of decomposition by poor C quality rather than from N inhibition of lignin decay. Even though ANPP is limited by N, decomposers are strongly limited by C quality. My results suggest that anthropogenic N deposition may increase leaf litter decomposition more in ecosystems characterized by low-lignin litter than in those characterized by high-lignin litter. Received 26 October 1999; accepted 2 June 2000.     

中国森林生态系统凋落叶分解速率的分布特征及其控制因子

凋落物分解是森林生态系统碳循环的重要组成部分。建立中国森林凋落叶分解速率数据库, 分析凋落叶分解速率与其主要影响因素之间的关系, 对精确地预测中国森林生态系统碳收支具有重要意义。该研究通过收集已报道的中国森林凋落叶分解常数(k)及其相关变量, 分析探讨地理因素(纬度、经度和海拔)、气候因素(年平均气温和年降水量)、凋落叶质量(氮、磷、钾、木质素、木质素:氮和碳氮比)和叶特性(常绿与落叶、阔叶与针叶)对中国森林凋落叶分解速率的影响。结果表明, 在国家尺度上, k随年平均气温、年降水量、氮、磷和钾的增加而增加, 随纬度、经度、海拔、碳氮比、木质素和木质素:氮的增大而减小, 叶特性对k的影响不显著。气候与地理因素(年平均气温、年降水量和纬度)能解释k值变异的34.1%, 凋落叶质量(氮、钾、木质素和木质素:氮)能解释k值变异的21.7%, 它们能共同解释k值变异的74.4%。了解森林凋落叶分解速率在国家尺度上的格局和主控因素可为中国森林生态系统碳循环相关模型提供基础参数。     

Extracellular Enzyme Activities and Carbon Chemistry as Drivers of Tropical Plant Litter Decomposition

Litter quality parameters such as nitrogen and lignin content correlate with decomposition rates at coarse scales, but fine‐scale mechanisms driving litter decomposition have proven more difficult to generalize. One potentially important driver of decomposition is the activity of extracellular enzymes that catalyze the degradation of complex compounds present in litter. To address the importance of this mechanism, we collected 15 Hawaiian plant litter types and decomposed them in fertilized and control plots for up to two years. We measured litter nutrient content and carbon chemistry prior to decomposition, as well as extracellular enzyme activities, mass loss, and litter nutrient content over time. We found that water‐soluble carbon content, cellobiohydrolase activities, and polyphenol oxidase activities were significantly correlated with mass loss. Enzyme activities and decomposition rate constants both varied significantly by litter type, and fertilization increased mass loss rates in five litter types. Some litter types that decayed faster under fertilization also showed time‐dependent increases in carbon‐degrading enzyme activities, but others decayed faster independent of enzyme changes. These results suggest that extracellular enzyme activities partially determine litter decomposition rates, but high soluble carbon content may circumvent the requirement for enzyme‐catalyzed decomposition.     

Spatial characteristics in decomposition rate of foliar litter and controlling factors in Chinese forest ecosystems

凋落物分解是森林生态系统碳循环的重要组成部分。建立中国森林凋落叶分解速率数据库, 分析凋落叶分解速率与其主要影响因素之间的关系, 对精确地预测中国森林生态系统碳收支具有重要意义。该研究通过收集已报道的中国森林凋落叶分解常数(k)及其相关变量, 分析探讨地理因素(纬度、经度和海拔)、气候因素(年平均气温和年降水量)、凋落叶质量(氮、磷、钾、木质素、木质素:氮和碳氮比)和叶特性(常绿与落叶、阔叶与针叶)对中国森林凋落叶分解速率的影响。结果表明, 在国家尺度上, k随年平均气温、年降水量、氮、磷和钾的增加而增加, 随纬度、经度、海拔、碳氮比、木质素和木质素:氮的增大而减小, 叶特性对k的影响不显著。气候与地理因素(年平均气温、年降水量和纬度)能解释k值变异的34.1%, 凋落叶质量(氮、钾、木质素和木质素:氮)能解释k值变异的21.7%, 它们能共同解释k值变异的74.4%。了解森林凋落叶分解速率在国家尺度上的格局和主控因素可为中国森林生态系统碳循环相关模型提供基础参数。     

Impact of fine litter chemistry on lignocellulolytic enzyme efficiency during decomposition of maize leaf and root in soil

Residue recalcitrance controls decomposition and soil organic matter turnover. We hypothesized that the complexity of the cell wall network regulates enzyme production, activity and access to polysaccharides. Enzyme efficiency, defined as the relationship between cumulative litter decomposition and enzyme activities over time, was used to relate these concepts. The impact of two contrasting types of cell walls on xylanase, cellulase and laccase efficiencies was assessed in relation to the corresponding changes in residue chemical composition (xylan, glucan, lignin) during a 43-day incubation period. The selected residues were maize roots, which are rich in secondary cell walls that contain lignin and covalent bridges between heteroxylans and lignin, and maize leaves having mostly non-lignified primary cell walls thus making the cellulose and hemicelluloses less resistant to enzymes. Relationships between C mineralization and change in residue quality through decomposition indicated that the level of substitution of arabinoxylans (arabinan to xylan ratio) provides a good explanation of the decomposition process. In leaves enriched in primary cell walls, arabinose substitution of xylan controlled C mineralization rate but hampered polysaccharide decomposition, but to a lesser extent than in roots in which arabinoxylans were mostly cross-linked with lignin. Enzyme activity was higher in leaf than root amended soils while enzyme efficiency was systematically higher in the presence of roots. This apparent paradox suggests that residue quality could preselect the microbial community. Indeed, we found that microorganisms exhibited an initial rapid growth in the presence of a high quality litter and produced enzymes that are not efficient in degrading recalcitrant cell walls while, in the presence of the more recalcitrant maize roots, microbial biomass grew more slowly but produced enzymes of higher efficiency. This high enzyme efficiency could be explained by the synergistic action of hydrolytic and oxidative enzymes even in the early stage of decomposition.     

Temporal Dynamics of Abiotic and Biotic Factors on Leaf Litter of Three Plant Species in Relation to Decomposition Rate along a Subalpine Elevation Gradient

Relationships between abiotic (soil temperature and number of freeze-thaw cycles) or biotic factors (chemical elements, microbial biomass, extracellular enzymes, and decomposer communities in litter) and litter decomposition rates were investigated over two years in subalpine forests close to the Qinghai-Tibet Plateau in China. Litterbags with senescent birch, fir, and spruce leaves were placed on the forest floor at 2,704 m, 3,023 m, 3,298 m, and 3,582 m elevation. Results showed that the decomposition rate positively correlated with soil mean temperature during the plant growing season, and with the number of soil freeze-thaw cycles during the winter. Concentrations of soluble nitrogen (N), phosphorus (P) and potassium (K) had positive effects but C:N and lignin:N ratios had negative effects on the decomposition rate (k), especially during the winter. Meanwhile, microbial biomass carbon (MBC), N (MBN), and P (MBP) were positively correlated with k values during the first growing season. These biotic factors accounted for 60.0% and 56.4% of the variation in decomposition rate during the winter and the growing season in the first year, respectively. Specifically, litter chemistry (C, N, P, K, lignin, C:N and lignin:N ratio) independently explained 29.6% and 13.3%, and the microbe-related factors (MBC, MBN, MBP, bacterial and fungal biomass, sucrase and ACP activity) explained 22.9% and 34.9% during the first winter and the first growing season, respectively. We conclude that frequent freeze-thaw cycles and litter chemical properties determine the winter decomposition while microbe-related factors play more important roles in determining decomposition in the subsequent growing season.     

Contrasting Effects of Substrate and Fertilizer Nitrogen on the Early Stages of Litter Decomposition

Commonly observed positive correlations between litter nitrogen (N) concentrations and decomposition rates suggest that N frequently limits decomposition in its early stages. However, numerous studies have found little, if any, effect of N fertilization on decomposition. I directly compared internal substrate N and externally supplied inorganic N effects on decomposition in sites varying in soil N availability. I decomposed eight substrates (with initial %N from 0–2.5) in control and N-fertilized plots at eight grassland and forest sites in central Minnesota. N fertilization increased decomposition at only two of eight sites, even though decomposition was positively related to litter N at all sites and to soil N availability across sites. The effect of externally supplied N on decomposition was independent of litter N concentration, but was greater at sites with low N availability. The inconsistent effects of substrate and externally supplied N may have arisen because decomposers use organic N preferentially as an N source; because inorganic N availability across sites or with fertilization induced changes in microbial community attributes (for example, lower C:N or greater efficiency) that reduced the response of decomposition to increased inorganic N supply; or because the positive correlation between litter N or site N availability with decomposition was spurious, caused by tight correlations between litter or site N and some other factor that truly limited decomposition. These inconsistent effects of substrate N and external N supply on decomposition suggest that the oft-observed relationship between litter N and decomposition may not indicate N limitation of decomposition.     

Response of litter decomposition and related soil enzyme activities to different forms of nitrogen fertilization in a subtropical forest

With the continuing increase in the impact of human activities on ecosystems, ecologists are increasingly becoming interested in understanding the effects of nitrogen deposition on litter decomposition. At present, numerous studies have investigated the effects of single form of nitrogen fertilization on litter decomposition in forest ecosystems. However, forms of N deposition vary, and changes in the relative importance of different forms of N deposition are expected in the future. Thus, identifying the effects of different forms of N deposition on litter decomposition in forest ecosystems is a pressing task. In this study, two dominant litter types were chosen from Zijin Mountain in China: Quercus acutissima leaves from a late succession broad-leaved forest and Pinus massoniana needles from an early succession coniferous forest. The litter samples were incubated in microcosms with original forest soil and treated with four different forms of nitrogen fertilization [NH₄ ⁺, NO₃ ⁻, CO(NH₂)₂, and a mix of all three]. During a 5-month incubation period, litter mass losses, soil pH values, and soil enzyme activities were determined. Results show that all four forms of nitrogen fertilization significantly accelerate litter decomposition rates in the broadleaf forest, while only two forms of nitrogen fertilization [i.e., mixed nitrogen and CO(NH₂)₂] significantly accelerate litter decomposition rates in the coniferous forest. Litter decomposition rates with the mixed nitrogen fertilization were higher than those in any single form of nitrogen fertilization. All forms of nitrogen fertilization enhanced soil enzyme activities (i.e., catalase, cellulase, invertase, polyphenol oxidase, nitrate reductase, urease, and acid phosphatase) during the litter decomposition process for the two forest types. Soil enzyme activities under the mixed nitrogen fertilization were higher than those under any single form of nitrogen fertilization. These results suggest that the type and activity of the major degradative enzymes involved in litter decomposition vary in different forest types under different forms of nitrogen fertilization. They also indicate that a long-term consequence of N deposition-induced acceleration of litter decomposition rates in subtropical forests may be the release of carbon stored belowground to the atmosphere.     

Factors influencing limit values for pine needle litter decomposition: a synthesis for boreal and temperate pine forest systems

We synthesized available data for decomposition of pine (Pinus) needle litter in pine forests to determine the litter chemical characteristics and climate factors that explained variation in the limit value, i.e. the level of accumulated mass loss at which the decomposition process either continues at a very low rate or possibly stops. Our data base included 56 separate studies on decomposition of pine needle litter, spanning Scots pine, lodgepole pine, Aleppo pine, stone pine and white pine, mainly incubated at the site of collection. Studies had 5 to 19 samplings, on average 10, and the decomposition was followed to a mass loss ranging from 47 to 83%, on average 67%. The periods from 3.0 to 5.4 years, on average 3.9 years, were of sufficient duration to allow estimates of limit values of decomposition. We used a linear mixed model with regression effects to relate limit values to potential explanatory variables, namely the sites’ long-term mean annual temperature (MAT) and mean annual precipitation (MAP) and to substrate-chemistry factors. Regarding the latter, we explored two models; one that included initial concentrations of water solubles, lignin, N, P, K, Ca, Mg, and Mn and one that included only lignin, N, Ca, and Mn to focus on those nutrients known to influence lignin degradation. Using backward elimination significant explanatory variables were determined. For litter decomposed in its site of origin we found the limit value to depend mainly on the initial concentration of Mn, with higher Mn concentrations resulting in higher accumulated mass loss. Thus, litter with higher Mn reached a higher limit value and left a smaller stable fraction. This is likely due to the fact that Mn is an essential component of ligninolytic enzymes important for degrading litter in the later stages of decomposition. Manganese has received little attention in decomposition studies to date. Given its significance in this synthesis, the role of Mn in influencing variation in the late stages of decomposition among ecosystems and among litters of other genera besides Pinus deserves further attention.     

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.     

The Influence of Nutrient Availability on Soil Organic Matter Turnover Estimated by Incubations and Radiocarbon Modeling

We investigated the decomposability of soil organic matter (SOM) along a chronosequence of rainforest sites in Hawaii that form a natural fertility gradient and at two long-term fertilization experiments. To estimate turnover times and pool sizes of organic matter, we used two independent methods: (1) long-term incubations and (2) a three-box soil model constrained by radiocarbon measurements. Turnover times of slow-pool SOM (the intermediate pool between active and passive pools) calculated from incubations ranged from 6 to 20 y in the O horizon and were roughly half as fast in the A horizon. The radiocarbon-based model yielded a similar pattern but slower turnover times. The calculation of the ¹⁴C turnover times is sensitive to the lag time between photosynthesis and incorporation of organic C into SOM in a given horizon. By either method, turnover times at the different sites varied two- or threefold in soils with the same climate and vegetation community. Turnover times were fastest at the sites of highest soil fertility and were correlated with litter decay rates and primary productivity. However, experimental fertilization at the two least-fertile sites had only a small and inconsistent effect on turnover, with N slowing turnover and P slightly speeding it at one site. These results support studies of litter decomposition in suggesting that while plant productivity can respond rapidly to nutrient additions, decomposition may respond much more slowly to added nutrients.     

Influence of Different Forest System Management Practices on Leaf Litter Decomposition Rates,Nutrient Dynamics and the Activity of Ligninolytic Enzymes: A Case Study from Central European Forests

Leaf litter decomposition is the key ecological process that determines the sustainability of managed forest ecosystems, however very few studies hitherto have investigated this process with respect to silvicultural management practices. The aims of the present study were to investigate the effects of forest management practices on leaf litter decomposition rates, nutrient dynamics (C, N, Mg, K, Ca, P) and the activity of ligninolytic enzymes. We approached these questions using a 473 day long litterbag experiment. We found that age-class beech and spruce forests (high forest management intensity) had significantly higher decomposition rates and nutrient release (most nutrients) than unmanaged deciduous forest reserves (P<0.05). The site with near-to-nature forest management (low forest management intensity) exhibited no significant differences in litter decomposition rate, C release, lignin decomposition, and C/N, lignin/N and ligninolytic enzyme patterns compared to the unmanaged deciduous forest reserves, but most nutrient dynamics examined in this study were significantly faster under such near-to-nature forest management practices. Analyzing the activities of ligninolytic enzymes provided evidence that different forest system management practices affect litter decomposition by changing microbial enzyme activities, at least over the investigated time frame of 473 days (laccase, P<0.0001; manganese peroxidase (MnP), P = 0.0260). Our results also indicate that lignin decomposition is the rate limiting step in leaf litter decomposition and that MnP is one of the key oxidative enzymes of litter degradation. We demonstrate here that forest system management practices can significantly affect important ecological processes and services such as decomposition and nutrient cycling.     

川西高山林线交错带凋落叶分解初期转化酶特征

胞外酶对于有机质的降解具有重要的作用。在凋落物分解过程中,酶活性不仅受到凋落物种类或基质质量的影响,还受到环境因素的影响。转化酶催化蔗糖水解为葡萄糖和果糖,因此在凋落物分解早期,转化酶比降解难分解物质的酶具有更重要的作用。以川西高山林线交错带12种代表性凋落叶为研究对象,对林线交错带不同植被类型下的凋落叶转化酶活性以及物种和环境因子对转化酶活性的影响进行了研究。结果表明:同一植被类型下,12个物种转化酶活性具有极显著差异(P0.01)。物种、环境因子及其交互作用对转化酶活性有极显著的影响(P0.01)。初始纤维素含量与转化酶活性极显著正相关(P0.01)。初始木质素和总酚含量与转化酶活性极显著负相关(P0.01),能够共同解释转化酶活性变异的50.8%。不同植物生活型中,禾草类转化酶活性均为最高,这可能与禾草类较高的初始纤维素含量、较低的木质素和总酚含量有关。多元线性回归分析表明,凋落叶含水量能单独解释转化酶活性变量的62.1%,是环境因子中最重要的变量。从植被类型来看,大多数物种的转化酶活性在针叶林中均极显著高于高山草甸和灌丛(P0.01),这可能与针叶林中凋落叶的含水量最高且雪被最厚有关。历经一个雪被期分解后,凋落叶初始质量与环境因子的综合作用能够解释转化酶活性变异的79.1%,表明川西高山林线交错带凋落叶分解前期转化酶活性主要受初始木质素含量、总酚含量和含水量的调控。在全球气候变化情景下,凋落物水分含量的变化将会强烈的影响凋落叶分解前期的转化酶活性。     

Litter decomposition of three lignin-deficient mutants of Sorghum bicolor during spring thaw

Decomposition of plant litter during the freeze-thaw season has recently gained attention as having a significant role in nutrient cycling in many cold ecosystems. However, few studies have examined decomposition of crop remnants during the freeze-thaw season in an agronomic setting when microbial activity is presumably low. We examined decomposition of four cultivars of sorghum (Sorghum bicolor) leaves in a field in Southern Minnesota, USA using the litterbag method. Three of the four cultivars we examined expressed the brown midrib (bmr) mutation which have altered/reduced levels of lignin in their secondary cell walls compared to the wild-type (WT). Litter was buried in the fall and harvested during the spring thaw. After 160 d the bmr mutants lost 57–62% of their initial mass, compared to 51% in the WT. Mass loss agreed with presumed initial litter quality, as the bmr litter had higher initial N, and holocellulose:lignin and lower lignin, C:N and lignin:N values compared to the WT. The increased decomposition of the bmr cultivars appears to be related to increased loss of hemicellulose and holocellulose (cellulose+hemicellulose) or higher initial N concentrations. Alterations in cell-wall deposition in the bmr cultivars may increase accessibility of microbial cell-wall degrading enzymes that accelerate mass loss. Our results demonstrate that alterations in initial lignin chemistry may influence decomposition of sorghum litter in an agronomic setting.     

Interactions among plants,bacteria, and fungi reduce extracellular enzyme activities under long‐term N fertilization

Atmospheric nitrogen (N) deposition has enhanced soil carbon (C) stocks in temperate forests. Most research has posited that these soil C gains are driven primarily by shifts in fungal community composition with elevated N leading to declines in lignin degrading Basidiomycetes. Recent research, however, suggests that plants and soil microbes are dynamically intertwined, whereby plants send C subsidies to rhizosphere microbes to enhance enzyme production and the mobilization of N. Thus, under elevated N, trees may reduce belowground C allocation leading to cascading impacts on the ability of microbes to degrade soil organic matter through a shift in microbial species and/or a change in plant–microbe interactions. The objective of this study was to determine the extent to which couplings among plant, fungal, and bacterial responses to N fertilization alter the activity of enzymes that are the primary agents of soil decomposition. We measured fungal and bacterial community composition, root–microbial interactions, and extracellular enzyme activity in the rhizosphere, bulk, and organic horizon of soils sampled from a long‐term (>25 years), whole‐watershed, N fertilization experiment at the Fernow Experimental Forest in West Virginia, USA. We observed significant declines in plant C investment to fine root biomass (24.7%), root morphology, and arbuscular mycorrhizal (AM) colonization (55.9%). Moreover, we found that declines in extracellular enzyme activity were significantly correlated with a shift in bacterial community composition, but not fungal community composition. This bacterial community shift was also correlated with reduced AM fungal colonization indicating that declines in plant investment belowground drive the response of bacterial community structure and function to N fertilization. Collectively, we find that enzyme activity responses to N fertilization are not solely driven by fungi, but instead reflect a whole ecosystem response, whereby declines in the strength of belowground C investment to gain N cascade through the soil environment.     

亚热带不同树种凋落叶分解对氮添加的响应

为探究不同质量凋落物对氮(N)沉降的响应, 该研究采用尼龙网袋分解法, 在亚热带福建三明格氏栲(Castanopsis kawakamii)自然保护区的米槠(Castanopsis carlesii)天然林, 选取4种本区常见的具有不同初始化学性质的树种凋落叶进行模拟N沉降(N添加)分解实验(施N水平为对照0和50 kg·hm ^-2·a ^-1)。研究结果表明: 在2年的分解期内, 对照处理的各树种凋落叶的分解速率依次为观光木(Michelia odora, 0.557 a ^-1)、米槠(0.440 a ^-1)、台湾相思(Acacia confusa, 0.357 a ^-1)、杉木(Cunninghamia lanceolata, 0.354 a ^-1); N添加处理凋落叶分解速率依次为观光木(0.447 a ^-1)、米槠(0.354 a ^-1)、杉木(0.291 a ^-1)、台湾相思(0.230 a ^-1), 除杉木凋落叶外, N添加显著降低了其他3种凋落叶分解速率。N添加不仅使4种树木凋落叶分解过程中的N释放减慢, 同时还抑制凋落叶化学组成中木质素和纤维素的降解; N添加在凋落叶分解过程中总体上提高β-葡萄糖苷酶(βG)和酸性磷酸酶活性, 对纤维素水解酶的活性影响不一致, 而降低β-N-乙酰氨基葡萄糖苷酶活性和酚氧化酶活性。凋落叶分解速率与凋落叶中的碳获取酶(βG)活性以及其化学组分中的可萃取物含量极显著正相关, 与初始碳浓度、纤维素和木质素含量极显著负相关, 与初始N含量没有显著相关性。凋落物类型和N添加的交互作用虽未影响干质量损失速率, 但对木质素和纤维素的降解具有显著效应。综上所述, 化学组分比初始N含量能更好地预测凋落叶分解速率, 而N添加主要通过抑制分解木质素的氧化酶(如PHO)来降低凋落叶分解速率。     

Extracellular Enzyme Activities and Soil Organic Matter Dynamics for Northern Hardwood Forests receiving Simulated Nitrogen Deposition

Anthropogenic nitrogen enrichment alters decomposition processes that control the flux of carbon (C) and nitrogen (N) from soil organic matter (SOM) pools. To link N-driven changes in SOM to microbial responses, we measured the potential activity of several extracellular enzymes involved in SOM degradation at nine experimental sites located in northern Michigan. Each site has three treatment plots (ambient, +30 and +80 kg N ha⁻¹ y⁻¹). Litter and soil samples were collected on five dates over the third growing season of N treatment. Phenol oxidase, peroxidase and cellobiohydrolase activities showed significant responses to N additions. In the Acer saccharum–Tilia americana ecosystem, oxidative activity was 38% higher in the litter horizon of high N treatment plots, relative to ambient plots, while oxidative activity in mineral soil showed little change. In the A. saccharum–Quercus rubra and Q. velutina–Q. alba ecosystems, oxidative activities declined in both litter (15 and 23%, respectively) and soil (29 and 38%, respectively) in response to high N treatment while cellobiohydrolase activity increased (6 and 39% for litter, 29 and 18% for soil, respectively). Over 3 years, SOM content in the high N plots has decreased in the Acer–Tilia ecosystem and increased in the two Quercus ecosystems, relative to ambient plots. For all three ecosystems, differences in SOM content in relation to N treatment were directly related (r² = 0.92) to an enzyme activity factor that included both oxidative and hydrolytic enzyme responses.     

Response of degradative enzymes to N fertilization during litter decomposition in a subtropical forest through a microcosm experiment

With the continuing increase in human activities causing accelerating rates of anthropogenic nitrogen deposition inputs into forests, there is considerable interest in understanding the effects of nitrogen deposition on litter decomposition. Two dominant litters were chosen from Zijin Mountain in China: Quercus acutissima from a broad-leaved forest and Pinus massoniana from a coniferous forest. The litters were incubated in microcosms and treated with a gradient of nitrogen fertilization. During a 6-month incubation, changes in chemical composition (i.e., lignin, total carbohydrate, and nitrogen), litter mass losses, soil pH values, and the activities of degradative enzymes were determined. Results showed that medium-nitrogen and high-nitrogen fertilization significantly accelerated litter decomposition rates of leaves, while only the high-nitrogen fertilization significantly accelerated litter decomposition rates of needles. The results also showed that cellulase and nitrate reductase were primarily responsible for litter decomposition in the broad-leaved forest, while catalase, cellulase, and acid phosphatase were primarily responsible for litter decomposition in the coniferous forest under conditions of no N fertilization; catalase, cellulase, and acid phosphatase were primarily responsible for litter decomposition in the broad-leaved forest, while catalase, cellulase, invertase, and nitrate reductase were primarily responsible for litter decomposition in the coniferous forest under conditions of N fertilization. Nitrogen fertilization-stimulated litter decomposition was due to the fact that the activities of enzymes, particularly cellulase, were accelerated.