森林土壤氮素转换及其对氮沉降的响应

近几十年人类活动向大气中排放的含氮化合物激增 ,并引起大气氮沉降也成比例增加。目前 ,氮沉降的增加使一些森林生态系统结构和功能发生改变 ,甚至衰退。近 2 0 a欧洲和北美有关氮沉降及其对森林生态系统的影响方面的研究较多 ,而我国少有涉及。森林土壤氮素转换是森林生态系统氮素循环的一个重要的组成部分 ,而矿化、硝化和反硝化作用是其核心过程 ,氮沉降作为驱动因子势必改变森林土壤氮素转换速度、方向和通量。根据国外近 2 0 a有关研究 ,首先介绍了森林土壤氮素转换过程和强度 ,论述森林土壤氮素在生态系统氮素循环中的作用 ,然后在此基础上 ,介绍了氮沉降对森林土壤氮素循环的研究途径 ,探讨了氮沉降对森林土壤氮素矿化、硝化和反硝化作用的影响及其机理     

森林土壤酶对环境变化的响应研究进展

全球气候变化已是不争的事实,对陆地生态系统特别是森林生态系统物质循环将产生显著的影响。土壤酶是森林土壤物质循环的主要限制因素之一,对气候变化的响应近年来受到广泛关注。由于森林土壤酶对全球气候变化的响应研究是预测未来环境变化对森林生态系统过程影响的关键,因此,着重综述了森林土壤酶对环境变化尤其是全球变暖和氮沉降响应方面的研究,并分析了未来研究的主要方向。环境变化会引起土壤p H、水分及其营养成分的变化,而这些变化会反作用于土壤酶的活性和稳定性。森林土壤酶对增温的响应,不仅与酶的种类以及增温的温度范围和持续时间有关,还与土壤类型有关,是多种因子综合作用的结果。森林土壤酶对氮添加的响应与林分类型和土层类型有关,受复合氮的影响更大。建议未来的研究应加强酶的基本性质对环境变化的响应研究,注重林分类型、土层类型导致的差异,强化多因素的交互作用,并进行长期、综合的观测。     

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。     

Control of Nitrogen Loss from Forested Watersheds by Soil Carbon:Nitrogen Ratio andTree Species Composition

Leaching losses of nitrate from forests can have potentially serious consequences for soils and receiving waters. In this study, based on extensive sampling of forested watersheds in the Catskill Mountains of New York State, we examine the relationships among stream chemistry, the properties of the forest floor, and the tree species composition of watersheds. We report the first evidence from North America that nitrate export from forested watersheds is strongly influenced by the carbon:nitrogen (C:N) ratio of the watershed soils. We also show that variation in soil C:N ratio is associated with variation in tree species composition. This implies that N retention and release in forested watersheds is regulated at least in part by tree species composition and that changes in species composition caused by introduced pests, climate change, or forest management could affect the capacity of a forest ecosystem to retain atmospherically deposited N. Received 4 March 2002; Accepted 4 June 2002.     

The nitrogen status of Austrian forest ecosystems as influenced by atmospheric deposition,biomass harvesting and lateral organomass exchange

Measurements of the deposition rates of atmospheric trace constituents to forest ecosystems in Austria have shown that the deposition of plant utilizable nitrogen compounds is in the range from 12 kg N to more than 30 kg N ha^-1 a^-1. Locally, even higher deposition rates are encountered as a consequence of point sources or special deposition mechanisms such as fog interception, hoar frost formation, and accumulation in snow drifts. In order to place these values into perspective, they are compared with the nitrogen demand of past and present forest land use and with natural processes of nitrogen depletion and accumulation in forest ecosystems. During wind erosion of forest litter, woody material with a wide C/N-ratio remains on the windward side of ridges, while nutrient-rich material with a narrow C/N-ratio is deposited on the leeward side. As a result, total nitrogen storage in the forest soil as well as overall C/N-ratios change dramatically along a transect over a ridge, thus indicating a strong influence of litter C/N ratio on nitrogen retention in the forest soil. A study of nitrogen stores in the soil of beech ecosystems of the same yield class in the Vienna Woods showed a significant correlation of total N-content with base saturation. These results suggest that nitrogen storage capacity of forest soils may be managed by liming and tree species selection. As knowledge is still meagre, a special study on factors which determine nitrogen storage in forest soils is proposed within the FERN-programme.     

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。     

Controls on soil nitrification and stream nitrate export at two forested catchments

Dormant season inorganic nitrogen (N) leaching varies considerably among forested catchments with similar bedrock, forest cover and deposition history. Recent work has highlighted the importance of winter rain-on-snow (ROS) events as a source of winter nitrate (NO₃-N) export, but differences among streams are likely due to differences in baseflow NO₃-N concentrations, and thus soil N processes. The objective of this study was to investigate rates of N-mineralization and nitrification as well as their potential environmental controls throughout the year, but with particular focus on the winter season in south-central Ontario, Canada. Field incubations were utilized to assess differences in NO₃-N and ammonium production over time and across topographic positions in two catchments with contrasting patterns of N export. Rates of nitrification were similar to rates of total mineralization, and nitrification rates were significantly higher during the summer and spring compared with the winter and fall; however, winter nitrification was substantial, and ranged from 19 to 36 % of annual rates. Seasonal differences in nitrification were largely driven by temperature, soil moisture and inorganic N concentration in soil. Rain and melting snow infiltrated the soil during ROS events, which were associated with increased NO₃-N availability, particularly in well-drained soils, and ROS-induced increases in stream nitrate concentrations were largest at the catchment dominated by well-drained soil. Annual nitrification fluxes were almost two orders of magnitude greater than N deposition or NO₃-N leaching fluxes at either catchment. Similar rates of NO₃-N production within the two catchments suggest that consumption of NO₃-N within wet soils is responsible for the 10-fold difference in NO₃-N export between the two streams. Notably, these results suggest that consumption processes were important for reducing NO₃-N export even during winter ROS events.     

Modeling of nitrogen dynamics in an Austrian alpine forest ecosystem on calcareous soils: a scenario-based risk assessment under changing environmental conditions

We modeled the behavior of an Austrian alpine forest ecosystem on calcareous soils under changing climate and atmospheric nitrogen deposition scenarios. The change of nitrate leaching, emission rates of nitrogen compounds, and forest productivity were calculated using four process-oriented models for the periods 1998-2002 and 2048-2052. Each model reflects with high detail a segment of the ecosystem: PnET-N-DNDC (photosynthesis-evapotranspiration-nitrification-denitrification-decomposition; short-term nitrogen cycling), BROOK90 (water balance for small and homogenous forest watersheds), HYDRUS (water flux in complex and heterogenous soils), and PICUS v1.3 (forest productivity). The nitrogen balance model (NBM) combines the individual results into a comprehensive picture and extends the specific values beyond the limits of the individual models. The evaluation of the findings was outlined with TRACE, a model enabling a long-term prognosis of nitrogen cycling in annual time steps. Temperature increase and nitrogen input are influenced by various components and processes of the forest ecosystem. An increase of the temperature of 2.5 degrees C led to an enhancement of the N2O emission rates and affected the mineralization and the nitrification rates with the consequence of increased nitrate leaching into the subsoil. Enhanced nitrogen input also showed notable effects on nitrate leaching.     

Temporal changes and fluxes of sulphur and calcium in wet and dry deposition, internal circulation as well as in run-off and soil in a forest at Gårdsjön, Sweden

Sulphur dioxide in air as well as dry deposition of sulphur dioxide to a forest has decreased by a factor of 20 during the last two decades. It was earlier found that the internal circulation of calcium in Norway spruce follows the dry deposition of sulphur dioxide. The sulphur and calcium fluxes from 1992 were calculated from wet deposition, throughfall and a surrogate surface. Earlier fluxes from 1981 to 1991 were calculated using assumptions of the dry deposition of non-marine sulphate and calcium. The new estimates confirm the earlier studies that the internal circulation of calcium in a coniferous forest is directly related to the dry deposition of sulphur dioxide to the canopies and that the internal circulation of calcium decreases at the same rate as the deposition of sulphur decreased during the last two decades. The deposition fluxes were also compared to run-off and uptake of calcium in the forest as well as on modelled weathering rates from severely acidified forested catchments near the coast and inland and the soil pool. A reconstruction of changes in the soil pool of calcium over the last 100 years indicate that the soil pool has decreased by ca. 70% in catchments with complete harvest of the forest and ca. 40% if branches and needles are left on the ground. In a natural forest without acid deposition the soil pool of calcium would have increased by 6%. Estimates for the 21st century show that harvesting of stems+branches and needles may almost empty the soil pool of calcium in the next 100 years. Increased nitrogen deposition has increased forest growth, which indirectly increased uptake of calcium by the trees and indirectly caused a further decrease of the soil pool of calcium.     

Nutrient supply of forest soils in relation to management and site history

Internal and external factors like forest management practices and atmospheric deposition may have large influences on the nutrient supply of forest soils. Examples are given for the effects of tree species, harvesting, site history, changes of species and specific soil conditions, nitrogen deposition, and forest growth dynamics. It is concluded from these examples that all of these factors may contribute to soil acidification in forest ecosystems under humid temperate climatic conditions.     

Ecological and site historical aspects of N dynamics and current N status in temperate forests

Ecological developments during Holocene age and high atmospheric depositions since industrialization have changed the N dynamics of temperate forest ecosystems. A number of different parameters are used to indicate whether the forests are N‐saturated or not, most common among them is the occurrence of nitrates in the seepage water below the rooting zone. The use of different definitions to describe N saturation implies that the N status of ecosystems is not always appropriately assessed. Data on N dynamics from 53 different German forests were used to classify various development states of forest ecosystems according to the forest ecosystem theory proposed by Ulrich for which N balances of input – (output plus plant N increment) were used. Those systems where N output equals N input minus plant N increment are described as (quasi‐) Steady State Type. Those forests where N output does not equal N input minus plant N increment as in a ‘transient state.’ Forests of the transient state may lose nitrogen from the soil (Degradation Type) or gain nitrogen [e.g., from atmospheric depositions (Accumulation Type)]. Forest ecosystems may occur in four different N states: (a) (quasi‐) Steady State Type with mull type humus, (b) Degradation Type with mull type humus, (c) Accumulation Type with moder type humus, and (d) (quasi‐) Steady State Type with moder type humus. Forests with the (quasi‐) steady state with mull type humus in the forest floor (n= 8) have high‐soil pH values, high N retention by plant increment, high N contents in the mineral soils, and have not undergone large changes in the N status. Forests of the Degradation Type lose nitrogen from the mineral soil (currently degradation is occurring on one site). Most forests that have moder or mor type humus and low‐soil pH values, and low N contents in the mineral soil have gone through the transient state of organic matter loss in the mineral soils. They accumulate organic matter in the forest floor (accumulation phase, currently 21 sites are accumulating 6–21 kg N ha^?1 yr^?1) or have reached a new (quasi‐) steady state with moder/mor type humus (n= 15). N retention in the accumulation phase has significantly increased in soil with N deposition (r²= 0.38), soil acidity (considering thickness of the forest floor as indices of soil acidity, r²= 0.43) and acid deposition (sulfate deposition, r²= 0.39). Retention of N (4–20 kg N ha^?1 yr^?1) by trees decreased and of soils increased with a decrease in the availability of base cations indicating the important role of trees for N retention in less acid soils and those of soils in more acid soils. Ecosystem theory could be successfully applied on the current data to understand the dynamics of N in temperate forest ecosystems.     

氮沉降对森林植物的影响

综述了氮沉降对森林植物的影响。氮沉降对森林植物的影响主要表现在以下6个方面：(1)在一定量范围内的氮沉降有利于植物的光合作用,但过量后则会引起植物的光合速率下降;(2)当植物生长受氮限制时,在一定程度上的氮沉降增加植物生产力,但当氮过量后,氮沉降则使植物的生产力下降;(3)过量的氮沉降导致植物体各种营养元素含量的比例失衡;(4)氮沉降会改变植物的形态结构,集中表现为根／冠比减小;(5)氮沉降会增加植物对天然胁迫如干旱、病虫害和风的敏感性,减少其抵御能力;(6)氮沉降会改变植物组成和降低森林植物的多样性。     

Combined global change effects on ecosystem processes in nine U.S. topographically complex areas

Concurrent changes in climate, atmospheric nitrogen (N) deposition, and increasing levels of atmospheric carbon dioxide (CO₂) affect ecosystems in complex ways. The DayCent-Chem model was used to investigate the combined effects of these human-caused drivers of change over the period 1980–2075 at seven forested montane and two alpine watersheds in the United States. Net ecosystem production (NEP) increased linearly with increasing N deposition for six out of seven forested watersheds; warming directly increased NEP at only two of these sites. Warming reduced soil organic carbon storage at all sites by increasing heterotrophic respiration. At most sites, warming together with high N deposition increased nitrous oxide (N₂O) emissions enough to negate the greenhouse benefit of soil carbon sequestration alone, though there was a net greenhouse gas sink across nearly all sites mainly due to the effect of CO₂ fertilization and associated sequestration by plants. Over the simulation period, an increase in atmospheric CO₂ from 350 to 600 ppm was the main driver of change in net ecosystem greenhouse gas sequestration at all forested sites and one of two alpine sites, but an additional increase in CO₂ from 600 to 760 ppm produced smaller effects. Warming either increased or decreased net greenhouse gas sequestration, depending on the site. The N contribution to net ecosystem greenhouse gas sequestration averaged across forest sites was only 5–7 % and was negligible for the alpine. Stream nitrate (NO₃ ^?) fluxes increased sharply with N-loading, primarily at three watersheds where initial N deposition values were high relative to terrestrial N uptake capacity. The simulated results displayed fewer synergistic responses to warming, N-loading, and CO₂ fertilization than expected. Overall, simulations with DayCent-Chem suggest individual site characteristics and historical patterns of N deposition are important determinants of forest or alpine ecosystem responses to global change.     

氮沉降下西南山地针叶林根际和非根际土壤微生物养分限制特征差异

长期氮(N)沉降诱导了土壤养分失衡, 深刻影响着森林生态系统养分循环过程、生态功能及其可持续发展。前期研究发现N沉降下西南森林树木生长受到不同程度的磷(P)限制, 而土壤微生物是否表现出与植物养分限制特征协同的响应仍未明确。基于此, 该研究以西南山地典型人工针叶林——华山松(Pinus armandii)林为对象, 通过野外原位模拟N沉降实验, 测定了土壤有效养分供给、土壤微生物生物量(碳(C)、N、P)含量以及胞外酶活性, 结合生态酶化学计量的3种模型(比值模型、矢量分析模型与阈值元素比率模型)验证森林根际/非根际土壤中微生物是否受P养分限制。结果表明: (1) N添加下两个土壤位置(根际和非根际土壤)酸性磷酸酶(AP)活性分别显著升高52.5%和53.2%, 导致土壤酶活性N:P分别降低7.8%和4.8%; (2)矢量模型分析发现N添加下两个土壤位置的矢量角度均大于45°, 根际土壤和非根际土壤的矢量角度分别为52.2°和49.0°; (3) N添加下两个土壤位置C:P阈值(TER_C:P)显著降低, 导致C:P阈值与土壤有效C:P的比值(TER_C:P/Av_C:P)远小于1, 且根际土壤表现更明显。综上所述, 3个模型均表明N沉降加剧了土壤微生物代谢的P限制, 且根际土壤微生物P限制程度更强, 这与土壤和微生物养分含量及其化学计量特征密切相关。该研究结果可为全球气候变化下森林生态系统的适应性管理提供重要科学依据。     

Impacts of tropospheric ozone and climate change on net primary productivity and net carbon exchange of China's forest ecosystems

Aim We investigated how ozone pollution and climate change/variability have interactively affected net primary productivity (NPP) and net carbon exchange (NCE) across China's forest ecosystem in the past half century. Location Continental China. Methods Using the dynamic land ecosystem model (DLEM) in conjunction with 10‐km‐resolution gridded historical data sets (tropospheric O₃ concentrations, climate variability/change, and other environmental factors such as land‐cover/land‐use change (LCLUC), increasing CO₂ and nitrogen deposition), we conducted nine simulation experiments to: (1) investigate the temporo‐spatial patterns of NPP and NCE in China's forest ecosystems from 1961–2005; and (2) quantify the effects of tropospheric O₃ pollution alone or in combination with climate variability and other environmental stresses on forests' NPP and NCE. Results China's forests acted as a carbon sink during 1961–2005 as a result of the combined effects of O₃, climate, CO₂, nitrogen deposition and LCLUC. However, simulated results indicated that elevated O₃ caused a 7.7% decrease in national carbon storage, with O₃‐induced reductions in NCE (Pg C year^?1) ranging from 0.4–43.1% among different forest types. Sensitivity experiments showed that climate change was the dominant factor in controlling changes in temporo‐spatial patterns of annual NPP. The combined negative effects of O₃ pollution and climate change on NPP and NCE could be largely offset by the positive fertilization effects of nitrogen deposition and CO₂. Main conclusions In the future, tropospheric O₃ should be taken into account in order to fully understand the variations of carbon sequestration capacity of forests and assess the vulnerability of forest ecosystems to climate change and air pollution. Reducing air pollution in China is likely to increase the resilience of forests to climate change. This paper offers the first estimate of how prevention of air pollution can help to increase forest productivity and carbon sequestration in China's forested ecosystems.     

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.     

The effect of increasing salinity and forest mortality on soil nitrogen and phosphorus mineralization in tidal freshwater forested wetlands

Tidal freshwater wetlands are sensitive to sea level rise and increased salinity, although little information is known about the impact of salinification on nutrient biogeochemistry in tidal freshwater forested wetlands. We quantified soil nitrogen (N) and phosphorus (P) mineralization using seasonal in situ incubations of modified resin cores along spatial gradients of chronic salinification (from continuously freshwater tidal forest to salt impacted tidal forest to oligohaline marsh) and in hummocks and hollows of the continuously freshwater tidal forest along the blackwater Waccamaw River and alluvial Savannah River. Salinification increased rates of net N and P mineralization fluxes and turnover in tidal freshwater forested wetland soils, most likely through tree stress and senescence (for N) and conversion to oligohaline marsh (for P). Stimulation of N and P mineralization by chronic salinification was apparently unrelated to inputs of sulfate (for N and P) or direct effects of increased soil conductivity (for N). In addition, the tidal wetland soils of the alluvial river mineralized more P relative to N than the blackwater river. Finally, hummocks had much greater nitrification fluxes than hollows at the continuously freshwater tidal forested wetland sites. These findings add to knowledge of the responses of tidal freshwater ecosystems to sea level rise and salinification that is necessary to predict the consequences of state changes in coastal ecosystem structure and function due to global change, including potential impacts on estuarine eutrophication.     

大气氮沉降与森林生态系统的氮动态

由于人类活动的影响,若干年代以来大气氮沉降明显增加。在森林地区,大气氮沉降的空间变异性由林分的位置、结构和组成树种所决定。除降雨之外,干沉降和隐藏降水也是大气氮沉降的重要形式。     

The complete nitrogen cycle of an N-saturated spruce forest ecosystem

Long-term nitrogen deposition into forest ecosystems has turned many forests in Central Europe and North America from N-limited to N-saturated systems, with consequences for climate as well as air and groundwater quality. However, complete quantification of processes that convert the N deposited and contributed to ecosystem N cycling is scarce. In this study, we provide the first complete quantification of external and internal N fluxes in an old-growth spruce forest, the Höglwald, Bavaria, Germany, exposed to high chronic N deposition. In this forest, N cycling is dominated by high rates of mineralisation of soil organic matter, nitrification and immobilisation of ammonium and nitrate into microbial biomass. The amount of ammonium available is sufficient to cover the entire N demand of the spruce trees. The data demonstrate the existence of a highly dynamic internal N cycle within the soil, driven by growth and death of the microbial biomass, which turns over approximately seven times each year. Although input and output fluxes are of high environmental significance, they are low compared to the internal fluxes mediated by microbial activity.     

森林土壤氮素的转化与循环

森林土壤氮素转化与循环在森林生态系统功能中占有极其重要的意义。本文综述了森林土壤氮素转化与循环的研究历程和现状 ;介绍了凋落物的归还、施肥、大气沉降、自生固氮、氨化、硝化、反硝化、植物吸收、NH3 的挥发、NO3 -淋溶等土壤氮素输入、转化和输出的途径和过程 ;最后从研究目标、研究方法、研究对象和研究内容 4个方面归纳了森林土壤氮素转化与循环的发展趋势。