Ecological impacts of nitrogen deposition on terrestrial ecosystems: research progresses and prospects

Due to huge consumption of fossil fuels and chemical fertilizers, substantial amount of anthropogenic reactive nitrogen (N) has been released into the environment. Therefore, N deposition has gradually increased worldwide and become one of the most important issues of global change. China has been a N deposition hotspot, and N deposition is projected to last long duration, which poses serious threats to ecosystem stability and functionality. In this synthesis paper, we summarized the impacts of N deposition on aboveground vegetation, soil microorganisms and biogeochemical cycling of major elements (carbon, N and phosphorus) in terrestrial ecosystems by outlining the progresses in the research field during the past 40 years. Results indicate that the accumulation of reactive N compounds induced by N deposition alters the soil environment, ecological stoichiometric balance and species co-occurrence patterns, thereby changing biodiversity and ecosystem functions. The rates, forms and duration of N deposition and the homeostasis of biosystem together with abiotic environments determine the direction and extent of the ecosystem response to N deposition. Through analysing local and foreign studies in this research area, we explore the weaknesses of relevant research that are being conducted in China. To advance the basic research on and risk management of N deposition, we propose the establishment of a N deposition monitoring and research network across the country with consideration of different ecosystems to promote regional and global risk assessments. Future research should highlight the combined multiple factors with N deposition and conduct direct and in-depth mechanism studies.     

Gaseous nitrogen emmissions from undisturbed terrestrial ecosystems: An assessment of their impacts on local and global nitrogen budgets

There is increasing interest in the importance of nitrogen gas emissions from natural (non-agricultural) ecosystems with respect to local as well as global nitrogen budgets and with respect to the effects of nitrogen oxides on atmospheric ozone levels and global warming. The volatile forms of nitrogen of common interest are ammonia (NH₃), nitrous oxide, (N₂O), dinitrogen (N₂), and NO_x (principally NO + NO₂). It is often difficult to attribute emissions of these compounds from soils to a single process because they are produced by a variety of common biogeochemical mechanisms. Although environmental conditions in the soil often appear to favor nitrogen gas emissions, the potential nitrogen gas emission rate from undisturbed ecosystems is rarely approached. The best estimates to date suggest that nitrogen gas emission rates from undisturbed ecosystems typically range from > 1 to perhaps 10 or 20 kg N ha^-1 yr^-1. Under certain conditions, however, emission rates may be much higher. For example, excreta from animals in grasslands may elevate ammonia volatilization up to 100 kg N ha^-1 yr^-1 depending on grazer density; tidal input of nutrients to coastal wetlands may support denitrification rates of several hundred kg N ha^-1 yr^-1 . Excepting such cases, gaseous nitrogen losses are probably a small component of the local nitrogen budget in most undisturbed ecosystems. However, emissions from undisturbed soils are an important component of the global source strengths for (N₂O + N₂), N₂O and NO_x (50%, 21%, and 10% respectively). Emission rates of N₂O from natural ecosystems are higher than assumed previously by perhaps 10 times. Large-scale disturbance may have a stimulatory effect on nitrogen emission rates which could have important effects on global nitrogen budgets. There is a need for more sophisticated methods to account for natural temporal and spatial variations of emissions rates, to more accurately and precisely assess their global source strengths.     

外来植物对陆地生态系统氮循环的影响途径

外来植物的引种和入侵已成为一个全球性的重要问题.许多外来植物不仅可以改变陆地生态系统的氮输入、输出,而且还可以通过改变氮素吸收、再利用,凋落物质量,土壤环境,土壤生物等因子影响陆地生态系统内都氮循环.在概述陆地生态系统氮循环基础上,系统综述了外来植物对陆地生态系统氮循环的影响方式和途径以及可能造成的生态后果,并对将来研究方向进行了展望:应更多考虑外来植物影响机制的复杂性、不同养分元素循环的相互作用和新技术手段的应用.     

Biological nitrogen fixation: rates,patterns and ecological controls in terrestrial ecosystems

New techniques have identified a wide range of organisms with the capacity to carry out biological nitrogen fixation (BNF)—greatly expanding our appreciation of the diversity and ubiquity of N fixers—but our understanding of the rates and controls of BNF at ecosystem and global scales has not advanced at the same pace. Nevertheless, determining rates and controls of BNF is crucial to placing anthropogenic changes to the N cycle in context, and to understanding, predicting and managing many aspects of global environmental change. Here, we estimate terrestrial BNF for a pre-industrial world by combining information on N fluxes with ¹⁵N relative abundance data for terrestrial ecosystems. Our estimate is that pre-industrial N fixation was 58 (range of 40–100) Tg N fixed yr⁻¹; adding conservative assumptions for geological N reduces our best estimate to 44 Tg N yr⁻¹. This approach yields substantially lower estimates than most recent calculations; it suggests that the magnitude of human alternation of the N cycle is substantially larger than has been assumed.     

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。     

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

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

降水变化和氮沉降影响森林叶根凋落物分解研究进展

全球环境变化通过改变凋落物质量和产量、土壤生物以及非生物因子调控森林凋落物分解,从而对森林生态系统物质和能量循环产生重要的影响。就森林凋落物分解对当前我国面临降水格局变化和大气氮沉降增加的响应进行了回顾和系统的分析,发现降水格局改变如降水减少可能降低凋落物质量从而减缓凋落物分解,而氮沉降增加通常提高凋落物质量从而促进凋落物分解（间接效应）;降水格局改变通过调节土壤含水量和溶解氧含量进而影响微生物参与的分解过程,或通过改变可溶性组分的淋溶量来影响凋落物分解的物理过程,而氮沉降增加主要通过提高外源氮素的有效性从而促进或抑制微生物参与的分解过程（直接效应）。现有研究大多是基于地上凋落物（例如叶凋落物）来理解和量化森林凋落物分解速率与环境因子之间的关系。但目前对降水格局变化及其与大气氮沉降增加的交互作用如何影响森林地上和地下凋落物分解,以及潜在的微生物学机制仍然缺乏统一和清晰的认识。从土壤性质、凋落物质量、微生物群落结构和功能3个方面构建了环境变化对森林地上和地下凋落物分解的概念框架,并进一步阐述未来研究的重点方向：（1）亟需查明地上和地下凋落物分解的驱动机制;（2）探明降水格局变化和氮添加单因子及两因子交互作用对凋落物分解和养分释放的影响及其生物化学调控机理;（3）阐明微生物群落结构和功能对降水格局变化和氮添加单因子及两因子交互的响应机制。以期为深入探讨全球环境变化对森林凋落物分解的影响,以及环境胁迫下森林土壤"碳库"维持机制的解释提供科学依据。     

Anthropogenic nitrogen sources and relationships to riverine nitrogen export in the northeastern U.S.A.

Human activities have greatly altered the nitrogen (N) cycle, accelerating the rate of N fixation in landscapes and delivery of N to water bodies. To examine relationships between anthropogenic N inputs and riverine N export, we constructed budgets describing N inputs and losses for 16 catchments, which encompass a range of climatic variability and are major drainages to the coast of the North Atlantic Ocean along a latitudinal profile from Maine to Virginia. Using data from the early 1990's, we quantified inputs of N to each catchment from atmospheric deposition, application of nitrogenous fertilizers, biological nitrogen fixation, and import of N in agricultural products (food and feed). We compared these inputs with N losses from the system in riverine export.The importance of the relative sources varies widely by catchment and is related to land use. Net atmospheric deposition was the largest N source (>60%) to the forested basins of northern New England (e.g. Penobscot and Kennebec); net import of N in food was the largest source of N to the more populated regions of southern New England (e.g. Charles & Blackstone); and agricultural inputs were the dominant N sources in the Mid-Atlantic region (e.g. Schuylkill & Potomac). Over the combined area of the catchments, net atmospheric deposition was the largest single source input (31%), followed by net imports of N in food and feed (25%), fixation in agricultural lands (24%), fertilizer use (15%), and fixation in forests (5%). The combined effect of fertilizer use, fixation in crop lands, and animal feed imports makes agriculture the largest overall source of N. Riverine export of N is well correlated with N inputs, but it accounts for only a fraction (25%) of the total N inputs. This work provides an understanding of the sources of N in landscapes, and highlights how human activities impact N cycling in the northeast region.     

土壤有机碳和氮分解对温度变化的响应趋势与研究方法

总结了土壤中碳和氮贮量与温度的关系、土壤碳和氮分解对温度时空差异和直接加热升温的响应,以及土壤碳和氮分解对低温冻结及冻融循环的响应趋势,讨论了其研究方法的误差和不确定性,并对今后的研究提出了一些建议．气候变暖在短期内将使土壤碳和氮分解加速并引起CO2释放量增加,而长期过程中却并不一定会引起土壤碳和氮分解加速．合理解释不同研究结果的差异,除了需要系统分析土壤碳和氮分解对温度变化响应的机制外,还需要充分认识土壤碳和氮分解对温度变化响应的长期过程和短期过程的差异,以及研究方法、植被、土壤和气候等因素的影响．     

氮沉降对森林土壤磷循环的影响

磷是生物体必需的大量元素之一,也是许多生态系统的主要限制因子。近年来,大气氮沉降日益加剧,已对森林生态系统磷循环产生了不可忽视的影响。关于氮沉降对生态系统磷循环的影响已开展了一系列的研究,然而尚缺少对其整体的认识。因此,通过收集国内外已发表的相关文章,从以下五个方面综述氮沉降对森林生态系统土壤磷循环的影响及其机理:1)阐述了森林生态系统土壤磷循环的概念;2)介绍了氮沉降对森林土壤磷循环影响的研究方法,包括长期定位模拟氮沉降法、自然氮沉降梯度法和同位素示踪法等;3)概述了氮沉降对森林生态系统土壤磷循环的影响。目前的研究结论趋向于认为长期氮沉降使森林土壤磷循环速率加快。长期氮输入易于使土壤中可溶性磷向非活性磷酸盐库迁移而难以被利用。因此,为了满足需求,土壤磷酸酶活性将增加以加速有机磷的矿化,从而加速磷素在土壤-植物-微生物之间的周转。4)探讨了氮沉降影响森林土壤磷循环的机制。氮沉降可通过改变土壤有机质的性质、微生物群落组成、磷酸酶活性以及阳离子的流动性等途径影响森林土壤磷循环;5)指出了当前研究存在的问题及未来的研究方向。     

飓风和台风对沿海地区森林生态系统的影响

飓风和台风是影响热带和温带沿海区域的主要灾害性气候之一,飓风和台风对于森林生态系统的影响是生态学关注的课题。综述了飓风和台风登陆对于森林生态系统树木和林分的危害影响形式及主要影响因素,着重举例阐述了树种和森林类型是影响台风危害程度的一个重要因素。分析了目前国际上开展的关于飓风和台风登陆对于森林生态系统碳、氮循环的影响,结果表明飓风、台风干扰导致的森林凋落物输入量、凋落物分解速率以及森林碳储存量动态变化较为复杂,与森林类型、林分空间位置以及台风过后的时间段密切相关。飓风引起的森林受损的恢复途径和机理与树冠受损严重程度直接相关,并受到光和水分条件的影响,及时的开花、结果以及充足的土壤种子库对森林植被恢复具有促进作用。在景观和区域尺度量化飓风和台风对沿海地区森林生态系统的影响也日益引起关注,在这方面,整合气象数据、遥感数据和地面调查的模型模拟方法起到重要的作用。今后应加强对于我国东南沿海地区森林生态系统遭受台风影响损失的生态监测和长期定位研究,加强关于台风对于不同森林生态系统类型和不同树种的危害形式和危害程度的研究,以及台风对于森林生态系统碳、氮循环影响的研究,弥补我国在以上领域的空白。     

氮添加对中国陆地植被地上-地下生物量分配的影响

大气氮沉降水平持续升高导致的外源氮输入增加,强烈影响了陆地生态系统的碳循环。目前,已有大量报道证实了氮沉降升高对全球陆地植被固碳的积极影响。虽然之前大部分研究将这一结果归因于光合作用增强导致的地上生物量增加,但最近的研究发现长期氮添加对植物地下根系的影响也同样重要。归纳整理了181篇公开发表的我国野外模拟氮沉降试验结果,采用整合分析(Meta-analysis)方法,定量评估了氮添加对我国陆地植被地上-地下生物量分配的影响特征和不同生态系统类型及施氮方式之间的影响差异。通过分析地上-地下生物量分配对氮添加的响应差异来探究植被碳增益对长期大气氮沉降增加的潜在响应机制。结果表明,氮添加显著增强了我国陆地植被的光合作用及碳固存,且植物碳增益在不同生态系统类型及施氮制度间有所差异。植物叶片的氮含量显著增加,使得叶片碳氮比及凋落物碳氮比显著降低,但并未显著影响细根的碳氮比。氮添加总体上显著提高了植物的净光合速率,但降低了光合利用效率。地上生物量,凋落物产量和根生物量平均分别显著增加了38%,17%和18%,总体上植物地上部分对氮添加的响应程度比地下部分更高。然而,不同生态系统类型的地上-地下生物...     

Global change, soil biodiversity, and nitrogen cycling in terrestrial ecosystems: three case studies

The relative contribution of different soil organism groups to nutrient cycling has been quantified for a number of ecosystems. Some functions, particularly within the N-cycle, are carried out by very specific organisms. Others, including those of decomposition and nutrient release from organic inputs are, however, mediated by a diverse group of bacteria, protozoa, fungi and invertebrate animals. Many authors have hypothesized that there is a high degree of equivalence and flexibility in function within this decomposer community and thence a substantial extent of redundancy in species richness and resilience in functional capacity. Three case studies are presented to examine the relationship between soil biodiversity and nitrogen cycling under global change in ecosystem types from three latitudes, i.e. tundra, temperate grassland and tropical rainforest. In all three ecosystems evidence exists for the potential impact of global change factors (temperature change, CO₂ enrichment, land-use-change) on the composition and diversity of the soil community as well as on various aspects of the nitrogen and other cycles. There is, however, very little unequivocal evidence of direct causal linkage between species richness and nutrient cycling efficiency. Most of the changes detected are shifts in the influence of major functional groups of the soil biota (e.g. between microflora and fauna in decomposition). There seem to be few data, however, from which to judge the significance of changes in diversity within functional groups. Nonetheless the soil biota are hypothesized to be a sensitive link between plant detritus and the availability of nutrients to plant uptake. Any factors affecting the quantity or quality of plant detritus is likely to change this link. Rigorous experimentation on the relationships between soil species richness and the regulation or resilience of nutrient cycles under global change thus remains a high priority.     

Sensitivity of continental United States atmospheric budgets of oxidized and reduced nitrogen to dry deposition parametrizations

Reactive nitrogen (N_r) is removed by surface fluxes (air–surface exchange) and wet deposition. The chemistry and physics of the atmosphere result in a complicated system in which competing chemical sources and sinks exist and impact that removal. Therefore, uncertainties are best examined with complete regional chemical transport models that simulate these feedbacks. We analysed several uncertainties in regional air quality model resistance analogue representations of air–surface exchange for unidirectional and bi-directional fluxes and their effect on the continental N_r budget. Model sensitivity tests of key parameters in dry deposition formulations showed that uncertainty estimates of continental total nitrogen deposition are surprisingly small, 5 per cent or less, owing to feedbacks in the chemistry and rebalancing among removal pathways. The largest uncertainties (5%) occur with the change from a unidirectional to a bi-directional NH₃ formulation followed by uncertainties in bi-directional compensation points (1–4%) and unidirectional aerodynamic resistance (2%). Uncertainties have a greater effect at the local scale. Between unidirectional and bi-directional formulations, single grid cell changes can be up to 50 per cent, whereas 84 per cent of the cells have changes less than 30 per cent. For uncertainties within either formulation, single grid cell change can be up to 20 per cent, but for 90 per cent of the cells changes are less than 10 per cent.     

Effects of nitrogen: phosphorus supply ratios on nitrogen fixation in agricultural and pastoral ecosystems

An analysis of data compiled from the literature confirms a strong inverse relationship between annual rates of nitrogen fixation and the soil nitrogen content in agricultural and pastoral ecosystems. However, this inverse relationship is strongly modified by the rate of application of phosphorus fertilizer, which strongly influences the activities of both symbiotic and non-symbiotic nitrogen fixing organisms. In the case of symbiotic legumes, the response of N-fixation to N and P is in part a result of changes in legume dominance within the plant community. These results, as well as supporting data presented from a review of experiments on nitrogen fixation in a variety of other terrestrial and aquatic ecosystems, provide important support for the hypothesis that phosphorus availability is a key regulator of nitrogen biogeochemistry. Published as Paper No. 9950, Journal Series, Nebraska Agricultural Research Division, University of Nebraska, Lincoln, NE, USA.     

冬季升温对高山生态系统碳氮循环过程的影响

全球温度升高是目前面临的重要环境问题,但存在明显的季节差异性,即冬季升温幅度显著高于夏季的季节非对称性趋势,这在高纬度和高海拔地区更加显著。冬季升温会直接影响积雪覆盖与冰冻层厚度,并引起冻融交替循环的增加,而冬季植物处于休眠状态,这会直接影响土壤中有效氮的吸收与损失,引起土壤有效氮可利用性的变化。然而,关于冬季增温对后续生长季节植物活动、土壤碳氮循环过程的影响等方面的研究仍存在诸多不确定。综述了冬季升温对积雪覆盖与冻融交替循环改变对高山生态系统物质循环的影响,以及冬季升温对土壤碳氮循环、微生物与酶活性的影响,并由此引起的植物物候期、群落结构、生产与养分循环与凋落物分解等生理、生态过程方面的研究进展。在未来的研究中,应针对不同生态系统特点选择合适的冬季增温方式,加强非极地苔原地区关于冬季升温的研究,注重关注冬季升温对植物-土壤微生物之间反馈作用的影响,重点关注冬季升温对生态系统的延滞效应。     

Nitrification,denitrification, and related functional genes under elevated CO2: A meta-analysis in terrestrial ecosystems

Global change may have profound effects on soil nitrogen (N) cycling that can induce positive feedback to climate change through increased nitrous oxide (N₂O) emissions mediated by nitrification and denitrification. We conducted a meta-analysis of the effects of elevated CO₂ on nitrification and denitrification based on 879 observations from 58 publications and 46 independent elevated CO₂ experiments in terrestrial ecosystems. We investigated the effects of elevated CO₂ alone or combined with elevated temperature, increased precipitation, drought, and N addition. We assessed the response to elevated CO₂ of gross and potential nitrification, potential denitrification, and abundances of related functional genes (archaeal amoA, bacterial amoA, nirK, nirS, and nosZ). Elevated CO₂ increased potential nitrification (+28%) and the abundance of bacterial amoA functional gene (+62%) in cropland ecosystems. Elevated CO₂ increased potential denitrification when combined with N addition and higher precipitation (+116%). Elevated CO₂ also increased the abundance of nirK (+25%) and nirS (+27%) functional genes in terrestrial ecosystems and of nosZ (+32%) functional gene in cropland ecosystems. The increase in the abundance of nosZ under elevated CO₂ was larger at elevated temperature and high N (+62%). Four out of 14 two-way interactions tested between elevated CO₂ and elevated temperature, elevated CO₂ and increased precipitation, and elevated CO₂ and N addition were marginally significant and mostly synergistic. The effects of elevated CO₂ on potential nitrification and abundances of bacterial amoA and nirS functional genes increased with mean annual temperature and mean annual precipitation. Our meta-analysis thus suggests that warming and increased precipitation in large areas of the world could reinforce positive responses of nitrification and denitrification to elevated CO₂ and urges the need for more investigations in the tropical zone and on interactive effects among multiple global change factors, as we may largely underestimate the effects of global change on soil N₂O emissions.     

陆地生态系统植被氮磷化学计量研究进展

因化学功能的耦合和元素的不可替代性, 植物对N、P的需求和利用存在严格的比例。植物N、P化学计量在不同功能群、生长地区、生长季、器官之间以及环境梯度下存在明显的变化规律。多数研究从N、P浓度、N:P及N、P间异速指数等角度分析了植物化学计量变化规律, 并探讨其在全球范围内的具体数值。为增进对植物响应全球变化的理解, 该文综述了N、P化学计量的影响因素及其机理的最新研究进展, 并指出未来拟重点研究的方向。     

氮磷添加对树木生长和森林生产力影响的研究进展

人为活动所导致的氮、磷输入和大气氮、磷沉降使生态系统中的氮、磷可利用性大幅提高, 对陆地生态系统的碳循环过程产生了显著影响。树木生长和森林生产力在全球碳循环中发挥着重要作用, 它决定着陆地碳固存的大小和方向。目前, 在全球范围内开展了很多氮、磷添加调控树木生长和森林生产力的野外控制实验, 但是研究结果并不一致, 受到多种生物、环境和实验处理条件等因素的影响。该文从野外氮添加和磷添加实验的文献数量、实验数量及其全球空间分布三个方面概述了氮、磷添加对树木生长和森林生产力影响的研究现状, 并总结了氮、磷添加实验中树木生长和森林生产力的评估方法, 包括相对生长速率和绝对增长量。基于相关的研究结果, 阐述了氮、磷添加影响树木生长和森林生产力的调控因素及其潜在影响机制, 包括气候、树木径级与林龄、植物功能性状(共生菌根类型、树木固氮属性和保守性与获得性性状)、植物和微生物相互作用关系、区域养分沉降速率和实验处理条件等。最后, 基于当前的研究进行了系统总结, 并指出今后需要加强的几个方面的研究, 以期为后续研究提供参考: 树木生长响应氮、磷添加的生理学机制, 树木各部分生长对氮、磷添加响应的权衡与分配, 植物功能性状在调节与预测树木生长响应氮、磷添加中的作用, 树木之间的竞争关系如何调控氮、磷添加对树木生长的影响, 以及开展长期的和联网的氮、磷添加对树木生长和森林生产力影响的野外控制实验。