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...     

氮添加对森林植物磷含量的影响及其机制

氮沉降对森林生态系统磷循环产生了不可忽视的影响, 尤其是加剧了植物生长的磷限制, 从而使得氮沉降背景下植物磷含量变化备受关注。该文综述了氮添加对森林植物磷含量的影响, 认为氮添加通过促进土壤磷酸酶活性进而提高土壤有效磷含量, 有利于植物的磷吸收并增加植物磷含量。同时, 森林植物磷含量对氮添加的响应还受物种、生活型以及施氮时间长短等因素的影响。基于森林植物磷含量对氮添加响应的差异性, 该文进一步探讨氮富集背景下森林植物磷含量变化的可能机制: 1)外源氮输入通过改变土壤中有效磷含量从而对植物磷的来源产生影响; 2)通过影响植物的根系分泌物、菌根共生和根系形态结构等进而影响植物的磷吸收能力; 3)通过影响植物的磷养分再分配、磷养分重吸收对植物磷利用效率产生影响。综上所述, 外源氮输入使植物磷含量发生改变, 首要原因是土壤有效磷含量的改变, 其次是植物磷吸收能力和磷利用效率的改变起调控作用。     

植物磷获取机制及其对全球变化的响应

磷是植物生长的必需元素,而陆地生态系统普遍存在磷限制,全球变化可能会影响土壤磷循环过程,进一步加剧磷限制,探讨植物磷获取策略对科学预测生态系统生产力如何适应全球变化具有重要意义。该文通过收集和梳理相关文献,从4个方面综述植物的磷获取机制及其对全球变化的响应:1)植物的磷饥饿响应机制;2)植物的磷获取途径和策略;3)土壤微生物对植物磷吸收的影响; 4)植物磷吸收对全球变化(温度升高、氮沉降和降水变化)的响应及其机制。该综述有助于深入理解全球变化背景下植物适应低磷胁迫的机理,也可为养分资源管理实践提供理论依据。     

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。     

氮添加对森林植物磷含量的影响及其机制

氮沉降对森林生态系统磷循环产生了不可忽视的影响,尤其是加剧了植物生长的磷限制,从而使得氮沉降背景下植物磷含量变化备受关注。该文综述了氮添加对森林植物磷含量的影响,认为氮添加通过促进土壤磷酸酶活性进而提高土壤有效磷含量,有利于植物的磷吸收并增加植物磷含量。同时,森林植物磷含量对氮添加的响应还受物种、生活型以及施氮时间长短等因素的影响。基于森林植物磷含量对氮添加响应的差异性,该文进一步探讨氮富集背景下森林植物磷含量变化的可能机制:1)外源氮输入通过改变土壤中有效磷含量从而对植物磷的来源产生影响; 2)通过影响植物的根系分泌物、菌根共生和根系形态结构等进而影响植物的磷吸收能力;3)通过影响植物的磷养分再分配、磷养分重吸收对植物磷利用效率产生影响。综上所述,外源氮输入使植物磷含量发生改变,首要原因是土壤有效磷含量的改变,其次是植物磷吸收能力和磷利用效率的改变起调控作用。     

土壤呼吸对降雨变化和氮沉降交互作用响应的研究进展

土壤呼吸作为陆地生态系统碳循环的关键过程,对大气CO₂浓度变化有直接影响。研究其如何响应降雨变化、氮沉降增加等全球变化因子,成为近年全球变化领域的热点与难点。与土壤呼吸响应降雨变化或氮沉降增加单个因子相比,研究土壤呼吸对这两个因子交互作用的响应更接近真实的自然环境,可更准确地预估未来土壤碳排放的变化趋势。目前,相关研究涉及全球不同的陆地生态系统,从土壤、微生物和植物层面对其响应机理进行揭示。本文从土壤呼吸及其组分、相关的土壤性质、微生物及植物因素方面,较全面地梳理了不同陆地生态系统土壤呼吸响应降雨变化和氮沉降增加交互作用的研究进展,指出了现有研究中的不足及今后需加强的研究方向,以期为进一步揭示土壤呼吸对降雨变化和氮沉降增加交互作用的响应规律及机制提供参考。     

环境因子对森林净生态系统生产力的影响

净生态系统生产力(net ecosystem production,NEP)是生态系统净的碳积累速率,可以指示生态系统碳汇/碳源的状态,当NEP为正值时指征生态系统为碳汇,反之则为碳源。在全球环境变化背景下,NEP已作为生态系统碳循环的核心概念被深入研究。本文以NEP为出发点,综述了5个主要非生物环境因子(水分、温度、氮沉降、大气CO2浓度增加和时空尺度)对森林生态系统NEP的影响。从文献分析表明NEP受生态系统本身性质和各环境影响因子及其之间相互作用的调控。本文最后指出,合理运用Meta分析、生态学联网研究、设计和开展长期观测、多尺度与多因子的科学试验在未来研究中的重要性和必要性,相关研究的开展将有利于全面理解和正确评估环境因子对净生态系统生产力的影响,对研究和预测全球变化对陆地生态系统碳循环的影响具有重要意义。     

Nitrogen addition affects plant biomass allocation but not allometric relationships among different organs across the globe

全球尺度上氮添加影响植物生物量分配但不影响不同器官间的异速生长关系 生物量在不同器官间的分配是植物对环境变化响应的一个关键生态生理学过程。然而,在全球尺度上有关不同陆地生态系统植物生物量分配对氮沉降响应的认识还比较欠缺。本文通过整合分析333篇已发表文章的5474个观测值,基于“最优分配假说”和“异速分配假说”,评估了全球尺度上氮添加对植物生物量及其在不同器官间分配的影响。结果表明：(1)氮添加显著增加了整株植物或不同器官的生物量,降低了根冠比和根质量分数,但对叶质量分数和茎质量分数无显著影响;(2)氮添加对不同器官质量分数的影响受实验条件、植物功能性状、纬度、氮添加率等因子单独或交互作用的调控;(3)氮添加对生物量在不同器官间的异速分配率无显著影响,表明氮添加导致的根冠比和根质量分数减少是在异速分配模式下由整株植物生物量增加而致。虽然氮添加改变了地上和地下部分的质量分数,但不同器官生物量间异速分配模式的稳定性表明“异速分配假说”能更好地描述植物生物量分配对氮添加的响应规律。该研究结果将有助于深入认识氮沉降环境下植物生物量的分配规律,同时表明将生物量分配纳入有关氮沉降对陆地生态系统影响的预测模型中的重要性。     

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。     

A review on the effects of nitrogen and phosphorus addition on tree growth and productivity in forest ecosystems

Nitrogen (N) and phosphorus (P) inputs induced by anthropogenic activities and atmospheric N and P deposition have largely increased the availability of soil N and P in terrestrial ecosystems, which have considerably affected terrestrial carbon cycling processes. Tree growth and productivity in forest ecosystems play an important role in global carbon cycling, and determine the magnitude and direction of terrestrial carbon sequestration. Currently, a large number of field manipulation experiments have been conducted to investigate the effects of N and/or P addition on tree growth and forest productivity, but the results from these studies were inconsistent. Such inconsistent results might be affected by multiple factors, including biological, environmental and experimental variables. Here, we reviewed the present research status of the effects of N and P addition on tree growth and forest productivity in forest ecosystems based on three aspects, including the number of publications and experiments with field N and P addition, and the global distributions of these experiments. Then, we summarized the methods for assessing tree growth and forest productivity at ecosystem level in forest ecosystems, including relative growth rate and absolute increment. According to the related results, we reviewed the regulating factors that affect tree growth and productivity, and the potential mechanisms for such factors, including climate, tree size and stand age, plant functional traits (including type of tree-associated mycorrhizal fungi, N-fixation property of trees, and conservative and acquisitive functional traits), plant-microbe interaction, ambient nutrient (i.e., N and P) deposition rate, and experimental variables. Finally, we summarized the current studies, and pointed out five aspects that are urgently needed to provide further insights in future studies, including the physiological mechanism of how tree growth responds to N and P addition, the tradeoff and allocation among growth of various parts of tree under N and P addition, the role of plant functional traits in regulating and predicting the responses of tree growth to N and P addition, how the competition among trees regulates the responses of tree growth to N and P addition, and conducting long-term and coordinated distributed field experiments investigating the effects of N and P addition on tree growth and forest productivity at the global scale.     

Global‐scale impacts of nitrogen deposition on tree carbon sequestration in tropical,temperate, and boreal forests: A meta‐analysis

Elevated nitrogen (N) deposition may increase net primary productivity in N‐limited terrestrial ecosystems and thus enhance the terrestrial carbon (C) sink. To assess the magnitude of this N‐induced C sink, we performed a meta‐analysis on data from forest fertilization experiments to estimate N‐induced C sequestration in aboveground tree woody biomass, a stable C pool with long turnover times. Our results show that boreal and temperate forests responded strongly to N addition and sequestered on average an additional 14 and 13 kg C per kg N in aboveground woody biomass, respectively. Tropical forests, however, did not respond significantly to N addition. The common hypothesis that tropical forests do not respond to N because they are phosphorus‐limited could not be confirmed, as we found no significant response to phosphorus addition in tropical forests. Across climate zones, we found that young forests responded more strongly to N addition, which is important as many previous meta‐analyses of N addition experiments rely heavily on data from experiments on seedlings and young trees. Furthermore, the C–N response (defined as additional mass unit of C sequestered per additional mass unit of N addition) was affected by forest productivity, experimental N addition rate, and rate of ambient N deposition. The estimated C–N responses from our meta‐analysis were generally lower that those derived with stoichiometric scaling, dynamic global vegetation models, and forest growth inventories along N deposition gradients. We estimated N‐induced global C sequestration in tree aboveground woody biomass by multiplying the C–N responses obtained from the meta‐analysis with N deposition estimates per biome. We thus derived an N‐induced global C sink of about 177 (112–243) Tg C/year in aboveground and belowground woody biomass, which would account for about 12% of the forest biomass C sink (1,400 Tg C/year).     

矿质养分输入对森林生物固氮的影响

生物固氮是森林生态系统重要的氮素来源,并且在全球氮循环中占有重要的地位。近代以来,因人类活动加剧而导致氮沉降的增加以及其它矿质养分元素(如磷、钼、铁等)输入的改变已成为影响森林生态系统生物固氮的重要因素之一,并引起了学术界的普遍关注。综述了国内外关于森林生物固氮对矿质养分输入的响应及机理。主要内容包括:(1)森林生物固氮的概念及主要的测定方法;(2)矿质养分输入对森林生物固氮的影响。整体上讲,氮素输入抑制了森林生物固氮,磷和其他营养元素输入则表现为促进作用。氮和磷、磷和微量元素同时添加均提高了森林的固氮量;(3)矿质养分改变森林生物固氮的机理。包括生物作用机制(如改变地表层固氮菌的数量或群落丰度、改变结瘤植物的根瘤生物量和附生植物的丰度或盖度)和环境作用机制(如引起土壤酸化、改变碳源物质的含量);(4)探讨了矿质养分输入对森林生物固氮影响研究中所存在的问题,并对未来该领域的研究提出建议。     

Biodiversity Promotes Tree Growth during Succession in Subtropical Forest

Losses of plant species diversity can affect ecosystem functioning, with decreased primary productivity being the most frequently reported effect in experimental plant assemblages, including tree plantations. Less is known about the role of biodiversity in natural ecosystems, including forests, despite their importance for global biogeochemical cycling and climate. In general, experimental manipulations of tree diversity will take decades to yield final results. To date, biodiversity effects in natural forests therefore have only been reported from sample surveys or meta-analyses with plots not initially selected for diversity. We studied biomass and growth of subtropical forests stands in southeastern China. Taking advantage of variation in species recruitment during secondary succession, we adopted a comparative study design selecting forest plots to span a gradient in species richness. We repeatedly censored the stem diameter of two tree size cohorts, comprising 93 species belonging to 57 genera and 33 families. Tree size and growth were analyzed in dependence of species richness, the functional diversity of growth-related traits, and phylogenetic diversity, using both general linear and structural equation modeling. Successional age covaried with diversity, but differently so in the two size cohorts. Plot-level stem basal area and growth were positively related with species richness, while growth was negatively related to successional age. The productivity increase in species-rich, functionally and phylogenetically diverse plots was driven by both larger mean sizes and larger numbers of trees. The biodiversity effects we report exceed those from experimental studies, sample surveys and meta-analyses, suggesting that subtropical tree diversity is an important driver of forest productivity and re-growth after disturbance that supports the provision of ecological services by these ecosystems.     

季节性雪被变化对森林凋落物分解及土壤氮动态的影响

全球气候变化引发的雪被格局变化将深刻影响植被的凋落物分解、陆地生态系统的土壤养分循环等过程.森林是陆地生态系统的主体,在全球生物地球化学循环中起着不可替代的作用.本研究综述了季节性雪被变化对森林凋落物分解及土壤氮动态的影响.全球气候变化情景下季节性雪被表现出因地域而异的增加或减少的变化格局,一方面通过改变环境温湿度、凋落物质量、分解者动态等直接影响分解过程,另一方面通过改变森林群落结构、植被物候、土壤养分等间接地作用于凋落物分解.同时,季节性雪被通过影响氮富集作用、雪被下土壤温湿度、冻融循环、森林群落、雪下动物和微生物等相关因子而改变森林土壤氮循环.本领域未来应开展的研究是: 1) 全面考虑全球气候变化情景下季节性雪被格局的变异性,开展不同季节性雪被格局变化的模拟研究;2) 开展季节性雪被融雪水淋溶作用对森林凋落物分解和土壤氮动态的影响研究;3) 阐明不同生态系统和气候带中季节性雪被格局变化对森林凋落物分解过程和土壤氮动态的驱动机制研究;4) 量化季节性雪被变化对森林凋落物分解和土壤氮动态在雪被覆盖期的瞬时影响和无雪期的延续影响,为阐明和模型预测陆地生态系统生物地球化学循环对全球气候变化的响应提供理论基础和数据支持.     

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。     

Functional traits determine tree growth and ecosystem productivity of a tropical montane forest: Insights from a long‐term nutrient manipulation experiment

Trait‐response effects are critical to forecast community structure and biomass production in highly diverse tropical forests. Ecological theory and few observation studies indicate that trees with acquisitive functional traits would respond more strongly to higher resource availability than those with conservative traits. We assessed how long‐term tree growth in experimental nutrient addition plots (N, P, and N + P) varied as a function of morphological traits, tree size, and species identity. We also evaluated how trait‐based responses affected stand scale biomass production considering the community structure. We found that tree growth depended on interactions between functional traits and the type or combination of nutrients added. Common species with acquisitive functional traits responded more strongly to nutrient addition, mainly to N + P. Phosphorous enhanced the growth rates of species with acquisitive and conservative traits, had mostly positive effects on common species and neutral or negative effects in rare species. Moreover, trees receiving N + P grew faster irrespective of their initial size relative to trees in control or to trees in other treatment plots. Finally, species responses were highly idiosyncratic suggesting that community processes including competition and niche dimensionality may be altered under increased resource availability. We found no statistically significant effects of nutrient additions on aboveground biomass productivity because acquisitive species had a limited potential to increase their biomass, possibly due to their generally lower wood density. In contrast, P addition increased the growth rates of species characterized by more conservative resource strategies (with higher wood density) that were poorly represented in the plant community. We provide the first long‐term experimental evidence that trait‐based responses, community structure, and community processes modulate the effects of increased nutrient availability on biomass productivity in a tropical forest.     

A tree-ring perspective on the terrestrial carbon cycle

Tree-ring records can provide valuable information to advance our understanding of contemporary terrestrial carbon cycling and to reconstruct key metrics in the decades preceding monitoring data. The growing use of tree rings in carbon-cycle research is being facilitated by increasing recognition of reciprocal benefits among research communities. Yet, basic questions persist regarding what tree rings represent at the ecosystem level, how to optimally integrate them with other data streams, and what related challenges need to be overcome. It is also apparent that considerable unexplored potential exists for tree rings to refine assessments of terrestrial carbon cycling across a range of temporal and spatial domains. Here, we summarize recent advances and highlight promising paths of investigation with respect to (1) growth phenology, (2) forest productivity trends and variability, (3) CO₂ fertilization and water-use efficiency, (4) forest disturbances, and (5) comparisons between observational and computational forest productivity estimates. We encourage the integration of tree-ring data: with eddy-covariance measurements to investigate carbon allocation patterns and water-use efficiency; with remotely sensed observations to distinguish the timing of cambial growth and leaf phenology; and with forest inventories to develop continuous, annually-resolved and long-term carbon budgets. In addition, we note the potential of tree-ring records and derivatives thereof to help evaluate the performance of earth system models regarding the simulated magnitude and dynamics of forest carbon uptake, and inform these models about growth responses to (non-)climatic drivers. Such efforts are expected to improve our understanding of forest carbon cycling and place current developments into a long-term perspective.