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。     

Impacts of Soil Nitrogen and Carbon Additions on Forest Understory Communities with a Long Nitrogen Deposition History

Rates of nitrogen (N) deposition have been historically high throughout much of the northeastern United States; thus, understanding the legacy of these high N loads is important for maintaining forest productivity and resilience. Though many studies have documented plant invasions due to N deposition and associated impacts on ecosystems, less is known about whether invasive plants will continue to increase in dominance with further shifting nutrient regimes. Using soil N and carbon additions, we examined the impact of both increasing and decreasing soil N on native and invasive understory plant dynamics over 4 years in a northeastern deciduous forest with a long history of N deposition. Despite applying large quantities of N, we found no difference in soil nitrate (NO₃) or ammonium (NH₄ ⁺) pools in N addition plots over the course of the study. Indicative of the potential N saturation in these forest soils, resin-available NO₃ ^? and NH₄ ⁺ showed evidence that the added N was rapidly moving out of the soil in N addition plots. Accordingly, we also found that adding N to soil altered neither invasive nor native plant abundance, though adding N temporally increased invasive plant richness. Carbon additions decreased soil N availability seasonally, but did not alter the total percent cover of invasive or native plants. Rather than being suppressed by excess N availability, native plant species in this ecosystem are primarily inhibited by the invasive species, which now dominate this site. In conclusion, understory plant communities in this potentially N-saturated ecosystem may be buffered to future alterations in N availability.     

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。     

A Framework to Assess Biogeochemical Response to Ecosystem Disturbance Using Nutrient Partitioning Ratios

Disturbances affect almost all terrestrial ecosystems, but it has been difficult to identify general principles regarding these influences. To improve our understanding of the long-term consequences of disturbance on terrestrial ecosystems, we present a conceptual framework that analyzes disturbances by their biogeochemical impacts. We posit that the ratio of soil and plant nutrient stocks in mature ecosystems represents a characteristic site property. Focusing on nitrogen (N), we hypothesize that this partitioning ratio (soil N: plant N) will undergo a predictable trajectory after disturbance. We investigate the nature of this partitioning ratio with three approaches: (1) nutrient stock data from forested ecosystems in North America, (2) a process-based ecosystem model, and (3) conceptual shifts in site nutrient availability with altered disturbance frequency. Partitioning ratios could be applied to a variety of ecosystems and successional states, allowing for improved temporal scaling of disturbance events. The generally short-term empirical evidence for recovery trajectories of nutrient stocks and partitioning ratios suggests two areas for future research. First, we need to recognize and quantify how disturbance effects can be accreting or depleting, depending on whether their net effect is to increase or decrease ecosystem nutrient stocks. Second, we need to test how altered disturbance frequencies from the present state may be constructive or destructive in their effects on biogeochemical cycling and nutrient availability. Long-term studies, with repeated sampling of soils and vegetation, will be essential in further developing this framework of biogeochemical response to disturbance.     

氮素和水分添加对贝加尔针茅草原植物多样性及生物量的影响

为了解草原植物群落物种多样性和植物地上生物量对氮沉降增加和降水变化的响应,在内蒙古贝加尔针茅(Stipa baicalensis)草原,分别设置对照(N0)、1.5 g/m2(N15)、3.0 g/m2(N30)、5.0 g/m2(N50)、10.0 g/m2(N100)、15.0 g/m2(N150)、20.0g/m2(N200)和30.0 g/m2(N300)(不包括大气沉降的氮量)8个氮素(NH4NO3)添加梯度和模拟夏季增加降水100mm的水分添加交互试验,研究氮素和水分添加对草原群落植物物种多样性和几种常见植物地上生物量的影响。结果表明:(1)氮素和水分的添加降低了草原群落植物物种多样性,且氮素和水分有显著的互作效应。在水分添加的条件下,随着施氮水平的增加,群落植物物种多样性减小;在无水分添加的条件下,随着施氮水平的增加,群落植物物种多样性呈先增加后减小的"单峰"变化趋势。(2)不同植物对氮素和水分添加的响应不同,随着施氮水平的增加,羊草地上生物量显著增加;贝加尔针茅、羽茅、糙隐子草、寸草苔和冷蒿先增加后减少,呈单峰曲线;星毛委陵菜、牧马豆、扁蓄豆和线叶菊地上生物量则逐渐减少。而且氮素和水分对贝加尔针茅、羽茅、扁蓄豆地上生物量有显著的交互作用。     

Impacts of nitrogen deposition on terrestrial plant diversity: a meta-analysis in China

Aims With the global atmospheric nitrogen (N) deposition increasing, the effect of N deposition on terrestrial plant diversity has been widely studied. Some studies have reviewed the effects of N deposition on plant species diversity; however, all studies addressed the effects of N deposition on plant community focused on species richness in specific ecosystem. There is a need for a systematic meta-analysis covering multiple dimensions of plant diversity in multiple climate zones and ecosystems types. Our goal was to quantify changes in species richness, evenness and uncertainty in plant communities in response to N addition across different environmental and experimental contexts.     

Convergent responses of nitrogen and phosphorus resorption to nitrogen inputs in a semiarid grassland

Human activities have significantly altered nitrogen (N) availability in most terrestrial ecosystems, with consequences for community composition and ecosystem functioning. Although studies of how changes in N availability affect biodiversity and community composition are relatively common, much less remains known about the effects of N inputs on the coupled biogeochemical cycling of N and phosphorus (P), and still fewer data exist regarding how increased N inputs affect the internal cycling of these two elements in plants. Nutrient resorption is an important driver of plant nutrient economies and of the quality of litter plants produce. Accordingly, resorption patterns have marked ecological implications for plant population and community fitness, as well as for ecosystem nutrient cycling. In a semiarid grassland in northern China, we studied the effects of a wide range of N inputs on foliar nutrient resorption of two dominant grasses, Leymus chinensis and Stipa grandis. After 4 years of treatments, N and P availability in soil and N and P concentrations in green and senesced grass leaves increased with increasing rates of N addition. Foliar N and P resorption significantly decreased along the N addition gradient, implying a resorption‐mediated, positive plant–soil feedback induced by N inputs. Furthermore, N : P resorption ratios were negatively correlated with the rates of N addition, indicating the sensitivity of plant N and P stoichiometry to N inputs. Taken together, the results demonstrate that N additions accelerate ecosystem uptake and turnover of both N and P in the temperate steppe and that N and P cycles are coupled in dynamic ways. The convergence of N and P resorption in response to N inputs emphasizes the importance of nutrient resorption as a pathway by which plants and ecosystems adjust in the face of increasing N availability.     

Nitrogen limitation of production in a Californianannual grassland: The contribution of arbuscularmycorrhizae

Nutrient availability limits plant production acrossa wide range of terrestrial ecosystems. In this studyof a Californian annual grassland community, theinfluence of arbuscular mycorrhizal (AM) associationson plant nutrient acquisition was investigated usingfactorial combinations of nitrogen (N), phosphorus (P)and benomyl fungicide. N additions resulted in asignificant increase in shoot biomass demonstratingthat plant productivity in these soils was N-limited. The effect of P additions consistently depended onfungicide treatment. In the absence of benomyl, shootP accumulation was unaltered by P addition. In thepresence of benomyl, P addition significantlyincreased shoot P accumulation and was associated witha consistent trend towards enhanced shoot biomass.The induction of P-deficiency with benomylapplication suggests that effective P acquisition byAM may contribute to the strong N-limitation ofproduction observed in many terrestrial ecosystems.     

草地群落多样性和生态系统碳氮循环对氮输入的非线性响应及其机制

理解草地生态系统结构和功能对氮富集的响应及其机制有助于准确评估大气氮沉降等外源氮输入的生态效应。全球范围内建立的多水平氮添加实验为认识草地生态系统结构和功能对氮输入的非线性响应机制提供了有效途径。为了反映学术界基于多水平氮添加控制实验取得的主要研究进展,该文综述了草地群落多样性和生态系统碳氮循环过程对外源氮输入的非线性响应特征及其驱动机制。按照目前的研究,氮输入会导致草地植物物种多样性、功能多样性以及土壤细菌多样性下降,但真菌多样性的变化并不明显。地上和地下生产力对氮输入的响应趋势存在差异:地上生产力沿氮添加梯度呈“先上升后饱和”的变化规律,而根系生产量和根冠比呈下降趋势,根系周转速率则呈“先上升后下降”的单峰格局。不同碳分解过程对氮输入的响应也不尽相同:凋落物分解速率沿氮添加梯度表现出“指数衰减、线性增加或无显著变化”的多元响应,而土壤呼吸和CH₄吸收速率与施氮量的关系则以“低氮促进、高氮抑制”的单峰趋势为主。类似地,不同土壤碳组分对氮输入的响应存在差异:氮添加总体会导致草地土壤碳库和颗粒态有机碳含量增加,而矿物结合态碳含量随施氮量呈“增加、不变或下降”的多元响...     

氮有效性增加对细根解剖、形态特征和菌根侵染的影响

氮(N)有效性增加对森林生态系统结构和功能有重要影响.细根作为树木地下最为活跃的组分,其对N有效性增加的响应已成为森林生态学研究的热点.本文对N有效性增加条件下细根解剖构造、形态和菌根侵染的响应趋势及潜在机制进行了综述.N有效性增加导致细根皮层厚度、皮层层数和皮层细胞直径下降,外皮层上通道细胞数量减少,而中柱和木质部横截面积,以及导管(或管胞)直径、数量和壁厚度均增加,根解剖构造的响应与植物激素浓度变化有关;N有效性增加后菌根侵染率下降,比根长(单位根干质量的长度)在外生菌根树种中普遍下降,而在内生菌根树种中增加;根直径和组织密度的变化趋势则表现出较大的种间差异.这些个体根和树种水平上的结果对于理解森林生态系统水平上的碳和养分循环具有重要意义.最后,根据目前研究中存在的不足提出了今后的研究方向与问题.     

15N自然丰度法在陆地生态系统氮循环研究中的应用

随着氮沉降的不断增加以及人们对全球变化问题的日益关注, 稳定同位素技术在全球变化研究中得到广泛的应用。因为植物和土壤的氮同位素组成记录了氮循环影响因子的综合作用, 并且具有测量简单以及不受取样时间和空间限制的优点, 所以氮同位素自然丰度法被用于氮循环的研究中。该文从氮循环过程中植物和土壤的氮分馏入手, 总结国内外相关文献, 阐述了植物和土壤氮自然丰度在预测生态系统氮饱和和氮循环长期变化趋势中的应用; 总结了利用树轮δ ¹⁵N法研究氮循环过程中应该注意的事项以及目前尚未解决的问题。     

Nitrogen fixation ability explains leaf chemistry and arbuscular mycorrhizal responses to fertilization

Atmospheric nitrogen (N) and phosphorus (P) deposition rates are predicted to drastically increase in the coming decades. The ecosystem level consequences of these increases will depend on how plant tissue nutrient concentrations, stoichiometry and investment in nutrient uptake mechanisms such as arbuscular mycorrhizal fungi (AMF) change in response to increased nutrient availability, and how responses differ between plant functional types. Using a factorial nutrient addition experiment with seedlings of multiple N-fixing and non-N-fixing tree species, we examined whether leaf chemistry and AMF responses differ between these dominant woody plant functional groups of tropical savanna and dry forest ecosystems. We found that N-fixers have remarkably stable foliar chemistry that stays constant with external input of nutrients. Non-N-fixers responded to N and N + P addition by increasing both concentrations and total amounts of foliar N, but showed a corresponding decrease in P concentrations while total amounts of foliar P stayed constant, suggesting a ‘dilution’ of tissue P with increased N availability. Non-N-fixers also showed an increase in N:P ratios with N and N + P addition, probably driven by both an increase in N and a decrease in P concentrations. AMF colonization decreased with N + P addition in non-N-fixers and increased with N and N + P addition in N-fixers, suggesting differences in their nutrient acquisition roles in the two plant functional groups. Our results suggest that N-fixers and non-N-fixers can differ significantly in their responses to N and P deposition, with potential consequences for future nutrient and carbon cycling in savanna and dry forest ecosystems.     

中国农村居民生活氮素流动特征

人类生活消费是陆地生态系统氮素流动的主要驱动力。定量核算和评估农村居民生活消费氮产生(N_RUR)及其活性氮(Nr)排放特征，对农村氮的可持续管理和生态环境改善具有重要的指导意义。以中国为例(2000—2020年),建立了N_RUR的产生及其活性氮排放核算框架。结果表明：20年来N_RUR上升了36.7%,年均5.62 Tg/a,食物消费氮是最大的贡献源(43.2%),工业日用品和家庭生活燃料消费氮分别占31.5%和25.3%;Nr排放量占N_RUR的25.4%(年均1.43 Tg/a),其以年均1.3%速率下降；NH₃挥发是最大的Nr排放源(50.1%),其次为排入地表水的Nr(31.0%)、NO_x(15.8%)和N₂O(2.0%),排入地下水的Nr仅占1.1%。加大人类粪尿排泄物的处置能力，减少秸秆燃料的使用比例、优化农村居民生活能源消费结构对农村居民生活消费Nr减排至关重要。     

Multi‐nutrient vs. nitrogen‐only effects on carbon sequestration in grassland soils

Human activities have greatly increased the availability of biologically active forms of nutrients [e.g., nitrogen (N), phosphorous (P), potassium (K), magnesium (Mg)] in many soil ecosystems worldwide. Multi‐nutrient fertilization strongly increases plant productivity but may also alter the storage of carbon (C) in soil, which represents the largest terrestrial pool of organic C. Despite this issue is important from a global change perspective, key questions remain on how the single addition of N or the combination of N with other nutrients might affect C sequestration in human‐managed soils. Here, we use a 19‐year old nutrient addition experiment on a permanent grassland to test for nutrient‐induced effects on soil C sequestration. We show that combined NPKMg additions to permanent grassland have ‘constrained’ soil C sequestration to levels similar to unfertilized plots whereas the single addition of N significantly enhanced soil C stocks (N‐only fertilized soils store, on average, 11 t C ha^?1 more than unfertilized soils). These results were consistent across grazing and liming treatments suggesting that whilst multi‐nutrient additions increase plant productivity, soil C sequestration is increased by N‐only additions. The positive N‐only effect on soil C content was not related to changes in plant species diversity or to the functional composition of the plant community. N‐only fertilized grasslands show, however, increases in total root mass and the accumulation of organic matter detritus in topsoils. Finally, soils receiving any N addition (N only or N in combination with other nutrients) were associated with high N losses. Overall, our results demonstrate that nutrient fertilization remains an important global change driver of ecosystem functioning, which can strongly affect the long‐term sustainability of grassland soil ecosystems (e.g., soils ability to deliver multiple ecosystem services).     

草地植物生产力主要影响因素研究综述

草地是全球分布面积最大的陆地生态系统,植物初级生产力是反映草地功能的重要指标。从植物种多样性、资源有效性、放牧、退化草地恢复和气候变化等方面较系统综述了影响草地植物生产力的关键要素和驱动力。大量研究表明,植物多样性与生产力的关系尚未有一致的结论,依据试验地点、起始状态甚至度量指标不同而不同;特别是资源有效性调节着生产力水平并对植物多样性和生产力关系产生显著影响;放牧改变了植物群落特征和养分有效性进而影响生产力的形成过程,也改变了资源有效性-植物多样性-生产力之间的关系;对于退化生态系统,在退化草地恢复过程中植物与土壤资源有效性的互作效应对植物生产力的变化起着关键作用;而在未来气候变化特别是增温对植物生产力的影响因地点和生态系统的不同而异,但多数研究结果显示增温提高了草地植物生产力。与国外其它草地分布区相比,国内的相关研究不仅在数量上明显不足,更重要是欠缺机理上的深入研究。在放牧和未来气候变化背景下如何维持和提高草地生产力,如何加速退化草地生态系统的恢复,进而实现生态安全建设和经济社会协调发展,是我国当前急需解决的理论和实践问题。     

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。     

磷的生物有效性对山地生态系统的影响

磷(P)的生物有效性对山地生态系统的发育和稳定至关重要。由于大气CO_2浓度升高和N沉降增加,生态系统C、N和P的化学计量比失衡,P的生物有效性受到更多关注。近年来山地系统中P的研究不断深入,2004—2013年间ISI Web of Knowledge中相关研究论文几乎是此前近百年的3倍。总结了山地生态系统中P的生物有效性的特点及其对植物物种多样性和初级生产力的影响。山地生态系统P的生物有效性因垂直高差和地形梯度空间变异明显,快速物质运移和生物过程是控制山地生态系统P的生物有效性的关键因素。P的生物有效性可以影响山地生态系统物种多样性和初级生产力,其对初级生产力的限制存在于全球范围内的山地生态系统。当P的生物有效性发生改变时,山地生态系统的结构越复杂,其植物物种多样性和初级生产力的响应可能会越平缓。全球变化的重要驱动因子(如增温和N沉降增加)可以直接或间接地改变山地生态系统P的生物有效性,因此需要在山地生态系统中加强长期监测和养分控制实验,并结合新型P分析技术,以期认识山地生态系统P的生物有效性的现状、变化趋势和对生态系统的影响,从而为适应全球变化背景下山地生态系统养分状况的改变提供依据。     

Combined effects of nitrogen addition and litter manipulation on nutrient resorption of Leymus chinensis in a semi‐arid grassland of northern China

Plant growth in semi‐arid ecosystems is usually severely limited by soil nutrient availability. Alleviation of these resource stresses by fertiliser application and aboveground litter input may affect plant internal nutrient cycling in such regions. We conducted a 4‐year field experiment to investigate the effects of nitrogen (N) addition (10 g N·m^?2·year^?1) and plant litter manipulation on nutrient resorption of Leymus chinensis, the dominant native grass in a semi‐arid grassland in northern China. Although N addition had no clear effects on N and phosphorus (P) resorption efficiencies in leaves and culms, N fertilisation generally decreased leaf N resorption proficiency by 54%, culm N resorption proficiency by 65%. Moreover, N fertilisation increased leaf P resorption proficiency by 13%, culm P resorption proficiency by 20%. Under ambient or enriched N conditions, litter addition reduced N and P resorption proficiencies in both leaves and culms. The response of P resorption proficiency to litter manipulation was more sensitive than N resorption proficiency: P resorption proficiency in leaves and culms decreased strongly with increasing litter amount under both ambient and enriched N conditions. In contrast, N resorption proficiency was not significantly affected by litter addition, except for leaf N resorption proficiency under ambient N conditions. Furthermore, although litter addition caused a general decrease of leaf and culm nutrient resorption efficiencies under both ambient and enriched N conditions, litter addition effects on nutrient resorption efficiency were much weaker than the effects of litter addition on nutrient resorption proficiency. Taken together, our results show that leaf and non‐leaf organs of L. chinensis respond consistently to altered soil N availability. Our study confirms the strong effects of N addition on plant nutrient resorption processes and the potential role of aboveground litter, the most important natural fertiliser in terrestrial ecosystems, in influencing plant internal nutrient cycling.     

Interactive effects of soil nitrogen and water availability on leaf mass loss in a temperate steppe

Although the link between leaf mass loss and assessment of ecosystem nutrient use efficiency and plant nutrient resorption efficiency has received considerable attention in various ecosystems, there has been relatively little effort to assess plant leaf mass loss during senescence, especially for herbaceous species. We conducted experimental studies to assess leaf mass loss during senescence in five dominant herbaceous species and examined the effects of increasing nitrogen (N) and water availability on leaf mass loss of four species in a temperate steppe in northern China. We nondestructively estimated mature leaf mass based on leaf length and width. Leaf mass loss varied substantially among species, ranging from 20–50%. On average across all species, N and water addition increased leaf mass loss by 30% and 19%, respectively. N and water addition interacted to affect leaf mass loss, as water addition had a significant positive effect on leaf mass loss under enriched N conditions but showed no effect under ambient N levels. We conclude that leaf mass loss of herbaceous plants was considerable and can potentially be more pronounced with increasing N and water availability. It is notable that the responses of plant species to N and water addition were variable. We suggest that leaf mass loss during senescence should be given full consideration in assessing nutrient use and resorption efficiency in semi-arid areas.     

NEECF: a project of nutrient enrichment experiments in China's forests

Anthropogenic nitrogen (N) emissions to atmosphere have increased dramatically in China since 1980s, and this increase has aroused great concerns on its ecological impacts on terrestrial ecosystems. Previous studies have showed that terrestrial ecosystems in China are acting as a large carbon (C) sink, but its potential in the future remains largely uncertain. So far little work on the impacts of the N deposition on C sequestration in China's terrestrial ecosystems has been assessed at a national scale. Aiming to assess and predict how ecological processes especially the C cycling respond to the increasing N deposition in China's forests, recently researchers from Peking University and their partners have established a manipulation experimental network on the ecological effects of the N deposition: Nutrient Enrichment Experiments in China's Forests Project (NEECF). The NEECF comprises 10 experiments at 7 sites located from north to south China, covering major zonal forest vegetation in eastern China from boreal forest in Greater Khingan Mountains to tropical forests in Hainan Island. This paper introduces the framework of the NEECF project and its potential policy implications.