土壤可溶性有机氮研究进展

可溶性有机氮(SON)和无机氮是陆地生态系统氮循环过程中重要的氮素形态,互为“源”和“汇”。陆地生态系统中氮素和其他营养元素的矿化、固持、淋溶和植物吸收均与SON有密切的联系。SON在土壤物质循环和养分流动等动态过程中的作用越来越受到关注,已成为生态学、环境学、土壤学、水文学等研究领域的热点之一。本文综述了国内外对土壤SON的研究进展,包括SON的定义和测定、SON库容大小和组成、植物和微生物对SON的吸收利用、SON来源及其影响因素、SON在土壤中的转化运移和淋失等。综合国内外研究结果发现,土壤SON是一个复杂的多组分可溶性有机物的混合物,主要为难降解的物质(惰性成分),能快速矿化分解的物质(活性成分)占比较低。由于惰性成分和活性成分在周转速率上的差异,SON在生态系统氮循环中的地位不能完全通过SON的容量特征来反映。因此,为了更准确地反映SON在氮周转、氮吸收和氮流失中的作用,未来需要创新研究方法并对SON组分加以区分:研究SON在氮转化和氮吸收中的作用时,重点关注SON中的活性成分;研究SON在氮淋溶径流损失中的贡献时,则重点关注SON中的惰性成分。     

氮稳定同位素技术在陆地生态系统氮循环研究中的应用

在过去几十年中, 氮(N)稳定同位素技术的发展提高了人们对于陆地生态系统氮循环的认识。该文回顾了氮稳定同位素技术在研究生态系统氮循环中的历史, 综述了最近十多年来氮稳定同位素技术在陆地生态系统氮循环研究中的典型案例, 包括利用氮同位素自然丰度鉴定植物氮来源、指示生态系统氮状态和量化过程速率, 利用¹⁵N标记技术示踪氮的去向和再分布等。该文同时指出这些应用中存在的问题, 以及在陆地生态系统上氮稳定同位素技术今后研究的重点发展方向。     

陆地生态系统植物多样性对矿质元素输入的响应

人类活动的加剧改变了陆地生态系统矿质元素(如氮、磷、钾等)循环的速度和方向,并且对生态系统的结构和功能也产生重要影响。如今,矿质元素输入量的改变及其产生的后续效应对陆地生态系统生物多样性的影响备受学者们的关注。从4个方面综述了全球氮沉降背景下主要矿质元素输入的改变对陆地植物多样性的影响及其机理:1)矿质营养元素限制的概念、确定方法以及与植物多样性的耦合关系;2)概述了氮、磷、钾等主要矿质元素输入对陆地植物多样性的影响:主要表现为负面效应;3)探讨了矿质元素输入影响植物多样性的可能机制,包括生态系统水平上的机制(如竞争排斥、酸化铝毒、物种入侵、同质性假说,间接诱导机制等)和植物个体水平上的机制(如元素失衡和环境敏感性增加等);4)根据目前研究现状,指出了已有研究的局限性,分析了未来可能的研究方向和重点。     

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.     

土壤有效氮及其相关因素对植物细根的影响

细根(直径≤2mm)作为植物吸收水分和养分的主要器官之一,在陆地生态系统养分循环和能量流动中起重要作用。开展土壤有效氮变化对植物细根影响研究对于了解全球气候变化条件下的陆地生态系统养分循环具有重要意义。本文就相关研究进行了综述:1)土壤有效氮变化对植物细根生长、发育、寿命及呼吸的直接影响;2)土壤质地、温度、大气CO2浓度和氮沉积等相关因素对植物细根的影响。由于研究方法及物种间差异等的影响,研究结果不尽相同。今后,应在不同空间尺度上深入研究土壤有效氮对植物细根的影响,而植物细根-土壤-微生物三者间相互关系变化对土壤氮变化的潜在响应将可能成为今后研究的热点问题之一。     

氮稳定同位素技术在陆地生态系统氮循环研究中的应用

在过去几十年中,氮(N)稳定同位素技术的发展提高了人们对于陆地生态系统氮循环的认识。该文回顾了氮稳定同位素技术在研究生态系统氮循环中的历史,综述了最近十多年来氮稳定同位素技术在陆地生态系统氮循环研究中的典型案例,包括利用氮同位素自然丰度鉴定植物氮来源、指示生态系统氮状态和量化过程速率,利用~(15)N标记技术示踪氮的去向和再分布等。该文同时指出这些应用中存在的问题,以及在陆地生态系统上氮稳定同位素技术今后研究的重点发展方向。     

丛枝菌根真菌形态结构、物种多样性和群落组成对氮沉降响应研究进展

工业革命以来,人类活动输入到生态系统中的氮迅速增加,已突破地球所能承受的氮循环阈值。过量氮沉降会造成生物多样性丧失等一系列危害,严重影响生态系统结构和功能。丛枝菌根(AM)真菌能够与大约70%-80%的陆地植物种类形成共生关系,在宿主植物养分吸收、抵抗外界不良环境压力、群落动态和物种共存、生物地球化学循环等方面具有重要的作用。探究AM真菌对氮沉降的响应对认识和把握菌根真菌缓解氮沉降的负面后果,维持生态系统的结构和功能具有重要意义。本文综述了AM真菌的形态结构、物种多样性和群落组成等对氮沉降的响应机制。前人研究表明氮沉降通常降低AM真菌的根系定殖率,减少根外菌丝密度和土壤孢子密度,改变菌丝生长的时间动态;降低AM真菌多样性,改变AM真菌群落组成。氮沉降主要通过缓解植物氮限制、降低植物对菌根的依赖性、减少植物对菌根的碳分配、改变根系和土壤中菌根生物量比率、在植物根内维持稳定的菌根真菌组成作为应对未来扰动的“保险”、改变土壤资源有效性及土壤酸度等直接和间接途径影响AM真菌结构和功能。我们建议在未来研究中整合多组学手段、开展学科交叉,聚焦复杂的生物互作体系对氮沉降的响应机制,以及AM真菌对氮沉降响应的生态后果。     

陆地生态系统氮沉降增加的生态效应

 人类活动在全球范围内极大地改变着氮素从大气向陆地生态系统输入的方式和速率，人为固定的氮素正在不断积累，并对生态系统的结构和功 能产生显著影响。该文从以下几个方面综述了大气氮沉降增加对陆地生态系统的影响：1)氮输入增加可能影响植物生产力和生态系统碳蓄积能 力，生态系统响应的方向和程度取决于系统的初始氮状况(氮限制或氮饱和)以及当地的植被和土壤特征；2)持续氮输入有可能改变土壤氮循环 过程，降低土壤固持氮的能力，甚至导致土壤酸化、盐基离子损耗，进而影响到土壤有机碳的分解；3)高的氮沉降速率和持续氮输入都可能加 速含氮痕量气体的释放，但其影响程度受生态系统初始状态的影响(例如磷限制和氮限制)；4)氮沉降增加会影响生态系统的物种丰富度、植物 群落结构和动态，促进森林扩张，改变菌根真菌的物种多样性；5)持续氮输入带来的植物群落结构和植物生理特征的变化可能影响昆虫取食特 性，进而通过食物链改变生态系统的营养结构；6) 氮沉降增加对生态系统的影响并不是孤立存在的，它与CO₂浓度升高和O₃浓度变化有协同作 用，但难以从其协同效应中区分出各自的影响。最后，该文总结了我国的氮沉降研究现状，并对今后的研究前景提出了展望。     

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。     

杉木人工林土壤可溶性有机质及其与土壤养分的关系

通过在福建省来舟林场对不同栽植代数杉木人工林土壤可溶性有机碳(DOC)和氮(DON)及土壤养分的研究,其结果表明,随着杉木栽植代数的增加林地土壤DOC和DON含量逐渐下降,在0～10cm土层内第3代杉木林土壤DOC和DON含量分别是第1代杉木林的83.9%和87.1%、第2代杉木林的90.6%和96.9%,在10～20cm土层内第3代杉木林土壤DOC和DON含量分别是第1代杉木林的80.2%和81.5%、第2代杉木林的81.8%和90.0%。在不同林地和土层内土壤DOC含量之间的差异性达到了显著或极显著水平,而DON含量之间的差异性不显著。不同栽植代数杉木林土壤养分的变化趋势与DOM一致,随着杉木连栽,土壤养分含量呈下降趋势。在0～10cm土层内第3代杉木林土壤全氮、全钾、铵态氮和速效钾含量分别是第1代杉木林的83.1%、60.4%、68.1%和44.3%,是第2代杉木林的84.6%、68.5%、74.4%和58.7%;在10～20cm土层内第3代杉木林土壤全氮、全钾、铵态氮和速效钾含量分别是第1代杉木林的74.0%、53.4%、57.6%和54.6%,是第2代杉木林的94.8%、59.5%、74.3%和65.5%。经相关性分析,在各土层内土壤DOC和DON含量与土壤全氮、全钾、铵态氮和速效钾等土壤养分含量存在着不同程度的相关性。     

川西高山林线土壤活性碳、氮对短期增温的响应

随着温室效应的加剧,受低温限制的高山林线生态系统对全球气候变暖较为敏感,可能直接影响到植物的生长和土壤碳氮过程.本研究假设气候变暖会改变高山生态系统土壤活性碳氮含量,在四川省理县米亚罗高山生态系统定位站,采用开顶式模拟增温装置(OTC)模拟增温对土壤活性碳、氮的短期影响.分别于2017年4、7和10月,采集OTC以及对照样地(CK)内土壤有机层和矿质土壤层的原状土壤,测定土壤可溶性有机碳(DOC)、土壤微生物生物量碳(MBC)、土壤可溶性有机氮(DON)和土壤微生物生物量氮(MBN)含量.结果表明: 模拟增温使年均气温升高0.88 ℃,土壤有机层和矿质土壤层的年均温度分别提高0.48和0.23 ℃.模拟增温没有显著改变土壤有机质和含水量,但显著提高了矿质土壤层的pH值,同时显著降低了非生长季矿质土壤层的DOC、DON含量;季节变化对两个层次的DOC、DON和MBN含量有极显著影响,而MBC没有明显的季节动态;增温和季节交互作用对矿质土壤层的DOC和DON有显著影响.土壤有机层的MBC、MBN含量显著高于矿质土壤层.土壤活性碳、氮与土壤有机质和含水量呈极显著正相关,MBC、MBN与土壤pH呈极显著正相关,MBN与土壤温度呈显著负相关.     

氮素添加和CO2浓度升高对白羊草根际和非根际土壤水溶性有机碳、氮的影响

采用盆栽控制试验对黄土丘陵区白羊草在不同CO₂浓度(400和800 μmol·mol^-1)和施氮水平(0、2.5、5.0 g N·m^-2·a^-1)条件下根际和非根际土壤水溶性有机碳(DOC)和水溶性有机氮(DON)的变化特征进行研究.结果表明: CO₂浓度升高对白羊草根际和非根际土壤DOC、水溶性总氮(DTN)、DON、水溶性铵态氮(NH₄⁺-N)、水溶性硝态氮(NO₃^--N)含量均无显著影响.施氮显著提高了根际和非根际土壤DTN、NO₃^--N含量和根际土壤DON含量,显著降低了根际土壤DOC/DON.在各处理条件下,根际土壤DTN、NO₃^--N和DON含量均显著低于非根际土壤,根际土壤DOC/DON显著高于非根际土壤.短期CO₂浓度升高对黄土丘陵区土壤水溶性有机碳、氮含量无显著影响,而氮沉降的增加在一定程度上改善了土壤中水溶性氮素缺乏的状况,但并不足以满足植被对水溶性氮素的需求.     

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.     

Patterns in the Chemical Fractionation of Organic Nitrogen in Rocky Mountain Streams

The intraannual dynamics of particulate organic nitrogen (PON) and two fractions of dissolved organic nitrogen (DON) were investigated in two Rocky Mountain streams draining watersheds with low rates of N deposition. Organic nitrogen accounted for over 60% of the total annual nitrogen export and consisted mostly of DON. Nitrate peaked during winter months and declined considerably during the growing season (less than 10 µg/L) suggesting the importance of biotic uptake. Concentrations of PON, total DON, and two DON fractions (humic and non-humic) peaked during spring runoff and were positively related to discharge, indicating hydrologic influence. Total DON and its two fractions showed significant inverse relationships to nitrate, indicating that DON and nitrate followed different intraannual patterns. Despite its seasonal fluctuations in concentration, PON showed a consistent carbon–nitrogen (C:N) ratio suggesting that it was relatively uniform in composition. Fractionation studies indicated that DON was primarily of non-humic origin, whereas dissolved organic carbon (DOC) was mainly derived from humic sources. The two DON fractions differed from each other in seasonal patterns of concentration and C:N ratio. The proportion of humic DON increased during snowmelt, and there were diverging seasonal patterns in the C:N ratio of the two fractions implying variations in bioavailability. Although organic nitrogen is commonly treated as a single pool in ecological studies, our results indicated that DON consists of fractions that undergo large intraannual changes in proportions and chemical composition. Treatment of DON as a single pool may be misleading from the viewpoint of understanding ecosystem processes directly related to changes in its sources and biological reactivity.     

Sources of organic nitrogen,phosphorus and carbon in antarctic streams

Dissolved and particulate organic materials were analysed in 14 streamwaters of the McMurdo Sound region of Antarctica. These streams are fed by glacial meltwaters and pass through catchments largely devoid of terrestrial vegetation. Nonetheless they contained measurable amounts of organic material in both dissolved and particulate form. Most of the dissolved organic carbon (DOC) values lay in the range 1–3 g C m^–3. Higher values were recorded close to penguin rookeries on the coast. Dissolved organic nitrogen (DON) concentrations were generally two orders of magnitude less than DOC and in flowing waters with rich blue-green algal growth DON increased with distance downstream. Dissolved organic phosphorus levels were generally much lower than DON, but highly variable. Particulate organic carbon concentrations (both fine and coarse) were unexpectedly high. Five sources of organic matter were identified: birdlife (only near the coast), autochthonous algal production (especially important for DON), streambed soils (important at first flows), lacustrine and marine sediments, through which certain streams and glaciers cut, and the glacial ice, which received organic input from wind-blown particulates, snowfall and the underlying bedrock of sedimentary origin. Highest organic levels were recorded in the first melt down the glacier face, suggesting that winter deposition of organic materials may be especially important.     

Seasonal Variations of Dissolved Nitrogen and DOC:DON Ratios in an Intermittent Mediterranean Stream

Seasonal variations of dissolved inorganic nitrogen (DIN) (NO₃–N and NH₄–N) and dissolved organic nitrogen (DON) were determined in Fuirosos, an intermittent stream draining an unpolluted Mediterranean forested catchment (10.5 km²) in Catalonia (Spain). The influence of flow on streamwater concentrations and seasonal differences in quality and origin of dissolved organic matter, inferred from dissolved organic carbon to nitrogen ratios (DOC:DON ratios), were examined. During baseflow conditions, nitrate and ammonium had opposite behaviour, probably controlled by biological processes such as vegetation uptake and mineralization activity. DON concentrations did not have a seasonal trend. During storms, nitrate and DON increased by several times but discharge was not a good predictor of nutrient concentrations. DOC:DON ratios in streamwater were around 26, except during the months following drought when DOC:DON ratios ranged between 42 and 20 during baseflow and stormflow conditions, respectively. Annual N export during 2000–2001 was 70 kg km⁻¹ year⁻¹, of which 75% was delivered during stormflow. The relative contribution of nitrogen forms to the total annual export was 57, 35 and 8% as NO₃–N, DON and NH₄–N, respectively.     

Fate and Transport of Organic Nitrogen in Minimally Disturbed Montane Streams of Colorado,USA

In two montane watersheds that receive minimal deposition of atmospheric nitrogen, 15–71% of dissolved organic nitrogen (DON) was bioavailable in stream water over a 2-year period. Discharge-weighted concentrations of bulk DON were between 102 and 135 μg/l, and the C:N ratio differed substantially between humic and non-humic fractions of DON. Approximately 70% of DON export occurred during snowmelt, and 40% of that DON was biologically available to microbes in stream sediments. Concentrations of bioavailable DON in stream water were 2–16 times greater than dissolved inorganic nitrogen (DIN) during the growing season, and bioavailable DON was depleted within 2–14 days during experimental incubations. Uptake of DON was influenced by the concentration of inorganic N in stream water, the concentration of non-humic DON in stream water, and the C:N ratio of the non-humic fraction of dissolved organic matter (DOM). Uptake of DON declined logarithmically as the concentration of inorganic N in stream water increased. Experimental additions of inorganic N also caused a decline in uptake of DON and net production of DON when the C:N ratio of non-humic DOM was high. This study indicates that the relative and absolute amount of bioavailable DON can vary greatly within and across years due to interactions between the availability of inorganic nutrients and composition of DOM. DOM has the potential to be used biotically at a high rate in nitrogen-poor streams, and it may be generated by heterotrophic microbes when DIN and labile DOM with low relative nitrogen content become abundant.     

The Role of Dissolved Organic Carbon, Dissolved Organic Nitrogen, and Dissolved Inorganic Nitrogen in a Tropical Wet Forest Ecosystem

Although tropical wet forests play an important role in the global carbon (C) and nitrogen (N) cycles, little is known about the origin, composition, and fate of dissolved organic C (DOC) and N (DON) in these ecosystems. We quantified and characterized fluxes of DOC, DON, and dissolved inorganic N (DIN) in throughfall, litter leachate, and soil solution of an old-growth tropical wet forest to assess their contribution to C stabilization (DOC) and to N export (DON and DIN) from this ecosystem. We found that the forest canopy was a major source of DOC (232 kg C ha^–1 y^–1). Dissolved organic C fluxes decreased with soil depth from 277 kg C ha^–1 y^–1 below the litter layer to around 50 kg C kg C ha^–1 y^–1 between 0.75 and 3.5m depth. Laboratory experiments to quantify biodegradable DOC and DON and to estimate the DOC sorption capacity of the soil, combined with chemical analyses of DOC, revealed that sorption was the dominant process controlling the observed DOC profiles in the soil. This sorption of DOC by the soil matrix has probably led to large soil organic C stores, especially below the rooting zone. Dissolved N fluxes in all strata were dominated by mineral N (mainly NO₃⁻). The dominance of NO₃^– relative to the total amount nitrate of N leaching from the soil shows that NO₃^– is dominant not only in forest ecosystems receiving large anthropogenic nitrogen inputs but also in this old-growth forest ecosystem, which is not N-limited.