稳定性同位素技术与Keeling曲线法在陆地生态系统碳/水交换研究中的应用

稳定性同位素技术和Keeling曲线法是现代生态学研究的重要手段和方法之一。稳定性同位素能够整合生态系统复杂的生物学、生态学和生物地球化学过程在时间和空间尺度上对环境变化的响应。Keeling曲线法是以生物过程前后物质平衡理论为基础,将CO₂或H₂O的同位素组成(δD、δ¹³C或δ¹⁸O)与其对应浓度测量结合起来,将生态系统净碳通量区分为光合固定和呼吸释放通量,或将整个生态系统水分蒸散区分为植物蒸腾和土壤蒸发。在全球尺度上,稳定性同位素技术、Keeling曲线法与全球尺度陆地生态系统模型相结合,还可区分陆地生态系统和海洋生态系统对全球碳通量的贡献以及不同植被类型(C₃或C₄)在全球CO₂同化量中所占的比例。然而,生态系统的异质性使得稳定性同位素技术和Keeling曲线法从冠层尺度外推到生态系统、区域或全球尺度时存在有一定程度的不确定性。此外,取样时间、地点的选取也会影响最终的研究结果。尽管如此,随着分析手段的不断精确和研究方法的日趋完善,稳定性同位素技术和Keeling曲线法与其它测量方法(如微气象法)的有机结合将成为未来陆地生态系统碳/水交换研究的重要手段和方法之一。     

The use of stable isotopes to study ecosystem gas exchange

Stable isotopes are a powerful research tool in environmental sciences and their use in ecosystem research is increasing. In this review we introduce and discuss the relevant details underlying the use of carbon and oxygen isotopic compositions in ecosystem gas exchange research. The current use and potential developments of stable isotope measurements together with concentration and flux measurements of CO₂ and water vapor are emphasized. For these applications it is critical to know the isotopic identity of specific ecosystem components such as the isotopic composition of CO₂, organic matter, liquid water, and water vapor, as well as the associated isotopic fractionations, in the soil-plant- atmosphere system. Combining stable isotopes and concentration measurements is very effective through the use of ”Keeling plots.” This approach allows the identification of the isotopic composition and the contribution of ecosystem, or ecosystem components, to the exchange fluxes with the atmosphere. It also allows the estimation of net ecosystem discrimination and soil disequilibrium effects. Recent modifications of the Keeling plot approach permit examination of CO₂ recycling in ecosystems. Combining stable isotopes with dynamic flux measurements requires precision in isotopic sampling and analysis, which is currently at the limit of detection. Combined with the micrometeorological gradient approach (applicable mostly in grasslands and crop fields), stable isotope measurements allow separation of net CO₂ exchange into photosynthetic and soil respiration components, and the evapotranspiration flux into soil evaporation and leaf transpiration. Similar applications in conjunction with eddy correlation techniques (applicable to forests, in addition to grasslands and crop fields) are more demanding, but can potentially be applied in combination with the Keeling plot relationship. The advance and potential in using stable isotope measurements should make their use a standard component in the limited arsenal of ecosystem-scale research tools. Received: 8 July 1999 / Accepted: 10 January 2000     

Stable isotope views on ecosystem function: challenging or challenged?

Stable isotopes and their potential for detecting various and complex ecosystem processes are attracting an increasing number of scientists. Progress is challenging, particularly under global change scenarios, but some established views have been challenged. The IX meeting of the Spanish Association of Terrestrial Ecology (AAET, Úbeda, 18–22 October 2009) hosted a symposium on the ecology of stable isotopes where the linear mixing model approach of partitioning sinks and sources of carbon and water fluxes within an ecosystem was challenged, and new applications of stable isotopes for the study of plant interactions were evaluated. Discussion was also centred on the need for networks that monitor ecological processes using stable isotopes and key ideas for fostering future research with isotopes.     

The use of stable carbon isotopes to decipher global change effects on soil organic carbon: present status,limitations, and future prospects

Major global change factors, including carbon dioxide (CO₂) fertilization, warming, change in precipitation, nitrogen deposition, and land-use change have the potential to significantly affect future stocks of soil organic carbon (SOC). These factors, individually or by interacting with each other, can also trigger positive or negative feedback to the processes affecting the rate of SOC formation or loss. Despite rapid progress in the understanding of carbon (C) cycling processes in the last few decades, much uncertainty remains in our ability to precisely forecast potential changes in SOC stocks in the rapidly changing future world. Stable C isotopes have been extensively used in natural observational studies as well as in laboratory and field experiments that manipulate CO₂ concentration, temperature, moisture, nitrogen fertilization, and vegetation type to understand the complex interactions and feedbacks that result from changing climate, plants and their herbivores, as well as soil microorganisms. Newly developed tools such as compound-specific isotope analysis, nano-SIMS (secondary ion mass spectroscopy), and stable isotope probing (SIP) permit isotope tracing in a specific ecosystem pool into specific C compounds and processes, thus providing in-depth insights into many processes affecting C biogeochemistry. The recent availability of affordable and reliable field-deployable optical isotope monitoring devices has provided researchers with a new set of tools for continuously tracking the ¹³C-CO₂ fluxes at the ecosystem level, enabling deeper insights into C biogeochemistry under changing environmental conditions. Despite these great strides, there is a scarcity of review studies that have comprehensively examined the use of C isotopes in studying SOC responses under global change factors. This review highlights recent progress in understanding the effect of major global change factors on SOC fluxes and stocks using selected examples covering scales from plant rhizospheres to geographic regions. Moreover, we discuss the strengths and limitations of current approaches and recent scientific advancements to highlight the new prospects evolving from the exceptional temporal and spatial resolution of stable isotope analysis in studying how global change affects SOC. Finally, we suggest that studies using stable C isotopes are well-poised to focus on identifying how dominant SOC cycling processes respond to environment-specific limiting factors and any thresholds and tipping points that define those relationships.

     

Empirical and theoretical challenges in aboveground–belowground ecology

A growing body of evidence shows that aboveground and belowground communities and processes are intrinsically linked, and that feedbacks between these subsystems have important implications for community structure and ecosystem functioning. Almost all studies on this topic have been carried out from an empirical perspective and in specific ecological settings or contexts. Belowground interactions operate at different spatial and temporal scales. Due to the relatively low mobility and high survival of organisms in the soil, plants have longer lasting legacy effects belowground than aboveground. Our current challenge is to understand how aboveground–belowground biotic interactions operate across spatial and temporal scales, and how they depend on, as well as influence, the abiotic environment. Because empirical capacities are too limited to explore all possible combinations of interactions and environmental settings, we explore where and how they can be supported by theoretical approaches to develop testable predictions and to generalise empirical results. We review four key areas where a combined aboveground–belowground approach offers perspectives for enhancing ecological understanding, namely succession, agro-ecosystems, biological invasions and global change impacts on ecosystems. In plant succession, differences in scales between aboveground and belowground biota, as well as between species interactions and ecosystem processes, have important implications for the rate and direction of community change. Aboveground as well as belowground interactions either enhance or reduce rates of plant species replacement. Moreover, the outcomes of the interactions depend on abiotic conditions and plant life history characteristics, which may vary with successional position. We exemplify where translation of the current conceptual succession models into more predictive models can help targeting empirical studies and generalising their results. Then, we discuss how understanding succession may help to enhance managing arable crops, grasslands and invasive plants, as well as provide insights into the effects of global change on community re-organisation and ecosystem processes.     

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。     

典型林区水分氢氧稳定同位素在土壤-植物-大气连续体中的分布特征

土壤-植物-大气连续体(SPAC)中水循环是水文学和生态学研究的重要内容,氢氧稳定同位素在不同水体中组成特征的差异可以指示水分循环过程.本研究通过分析成都平原区亚热带常绿阔叶林中降水、土壤水、植物水的同位素组成,探讨SPAC系统中水分的氢氧稳定同位素演化特征,揭示区域水循环不同界面过程.结果 表明:研究区雨季大气降水线...     

A method for in situ monitoring of the isotope composition of tree xylem water using laser spectroscopy

Field studies analyzing the stable isotope composition of xylem water are providing important information on ecosystem water relations. However, the capacity of stable isotopes to characterize the functioning of plants in their environment has not been fully explored because of methodological constraints on the extent and resolution at which samples could be collected and analysed. Here, we introduce an in situ method offering the potential to continuously monitor the stable isotope composition of tree xylem water via its vapour phase using a commercial laser‐based isotope analyser and compact microporous probes installed into the xylem. Our technique enables efficient high‐frequency measurement with intervals of only a few minutes per sample while eliminating the need for costly and cumbersome destructive collection of plant material and laboratory‐based processing. We present field observations of xylem water hydrogen and oxygen isotope compositions obtained over several days including a labelled irrigation event and compare them against results from concurrent destructive sampling with cryogenic distillation and mass spectrometric analysis. The data demonstrate that temporal changes as well as spatial patterns of integration in xylem water isotope composition can be resolved through direct measurement. The new technique can therefore present a valuable tool to study the hydraulic architecture and water utilization of trees.     

稳定氢氧同位素定量植物水分来源的不确定性解析

稳定氢氧同位素技术被广泛运用于生态系统、特别是干旱区生态系统中植物水分来源的研究,其理论假设为"水分被植物根系吸收并向木质部运输过程中不发生氢氧同位素分馏"。生态系统中不同水源的氢氧同位素组成普遍存在显著差异,为从水源混合体中区分出各水源的贡献率提供了前提条件。但在实际应用过程中,诸多因素导致稳定氢氧同位素技术定量植物水分来源的结果具有不确定性。综合已有研究并加以分析,举证说明植物吸收水分相对于水源同位素变化的滞后性、水源同位素的季节性变化、蒸发作用和水源之间的混合作用对水源同位素的影响等导致植物水分来源定量结果不确定性的几个因素,以期为今后稳定氢氧同位素技术在植物水分来源领域的应用提供参考。     

稳定同位素技术在植物水分利用研究中的应用

近20a稳定同位素技术在植物生态学研究中的应用得到了长足发展,使得对植物与水分关系也有了更深一步的了解。介绍稳定同位素性碳、氢、氧同位素在研究植物水分关系中的应用及进展,以期能为国内植物水分利用研究提供参考。由于植物根系从土壤中吸收水分时并不发生同位素分馏,对木质部水分同位素分析有助于对植物利用水分来源,生态系统中植物对水分的竞争和利用策略的研究,更好地了解生态系统结构与功能。稳定碳同位素作为植物水分利用效率的一个间接指标,在不同水分梯度环境中,及植物不同代谢产物与水分关系中有着广泛的应用。同位素在土壤-植被-大气连续体水分中的应用,有助于了解生态系统的水分平衡。随着稳定同位素方法的使用,植物与水分关系的研究将取得更大的进展。     

稳定同位素技术在土壤跳虫研究上的应用

稳定同位素技术的发展和应用是20世纪90年代生态学研究方法最重要的进步之一.许多生态学过程都伴随着同位素比率的变化,根据这些变化可以追踪生态系统的物质循环和能量流动.近年来,许多学者把稳定同位素技术引入到土壤生态学,主要用于研究土壤碳循环和土壤生物之间的营养关系,在很大程度上提升了对地下生态系统的理解.跳虫作为土壤动物群落的重要组分,它的食性和营养位置一直存在着许多不确定性.稳定同位素技术的应用,为确定跳虫的食物来源、营养位置和营养关系提供了一个强大的工具.本文综述了稳定同位素技术在跳虫食性和营养级研究上的应用,并指出了不足之处以及今后的发展方向.     

植物与土壤微生物在调控生态系统养分循环中的作用

陆地生态系统的地上、地下是相互联系的。植物与土壤微生物作为陆地生态系统中的重要组成部分, 它们之间的相互作用是生态系统地上、地下结合的重要纽带。该文首先介绍了植物在养分循环中对营养元素的吸收、积累和归还等作用, 阐述了土壤微生物对养分有效性及土壤质量具有重要的作用。其次, 重点综述了植物与土壤微生物之间相互依存、相互竞争的关系。植物通过其凋落物与分泌物为土壤微生物提供营养, 土壤微生物作为分解者提供植物可吸收的营养元素, 比如共生体菌根真菌即可使植物根与土壤真菌达到互惠。然而, 植物的养分吸收与微生物的养分固持同时存在, 因而两者之间存在对养分的竞争。通过植物多样性对土壤微生物多样性的影响分析, 以及土壤微生物直接或间接作用于植物多样性和生产力的分析, 探讨了植物物种多样性与土壤微生物多样性之间的内在联系。针对当前植物与土壤微生物对养分循环的调控机制的争论, 提出植物凋落物是调节植物与土壤微生物养分循环的良好媒介, 植物与土壤微生物的共同作用对维持整个生态系统的稳定性具有重要意义。也指出了目前在陆地生态系统地上、地下研究中存在的不足和亟待解决的问题。     

Carbon isotopes in terrestrial ecosystem pools and CO2 fluxes

Stable carbon isotopes are used extensively to examine physiological, ecological, and biogeochemical processes related to ecosystem, regional, and global carbon cycles and provide information at a variety of temporal and spatial scales. Much is known about the processes that regulate the carbon isotopic composition (delta(13)C) of leaf, plant, and ecosystem carbon pools and of photosynthetic and respiratory carbon dioxide (CO(2)) fluxes. In this review, systematic patterns and mechanisms underlying variation in delta(13)C of plant and ecosystem carbon pools and fluxes are described. We examine the hypothesis that the delta(13)C of leaf biomass can be used as a reference point for other carbon pools and fluxes, which differ from the leaf in delta(13)C in a systematic fashion. Plant organs are typically enriched in (13)C relative to leaves, and most ecosystem pools and respiratory fluxes are enriched relative to sun leaves of dominant plants, with the notable exception of root respiration. Analysis of the chemical and isotopic composition of leaves and leaf respiration suggests that growth respiration has the potential to contribute substantially to the observed offset between the delta(13)C values of ecosystem respiration and the bulk leaf. We discuss the implications of systematic variations in delta(13)C of ecosystem pools and CO(2) fluxes for studies of carbon cycling within ecosystems, as well as for studies that use the delta(13)C of atmospheric CO(2) to diagnose changes in the terrestrial biosphere over annual to millennial time scales.     

Isotopic Fractionation of Hydrogen in Plants

The naturally-occurring stable isotopes deuterium and hydrogen are fractionated by a number of physical and biological processes. Deuterium has a tendency to precipitate out first from a moist air mass. Thus ground water will become isotopically lighter with an increase in latitude, altitude, or distance inland. Water taken up by the plant from the soil undergoes little change until evapotranspiration results in leaf water becoming isotopically heavier. Thus hydrogen isotopes in plants can reveal something of geography (groundwater) and climate. Hydrogen isotopes undergo little fractionation by passage through the food chain, although plant parasites tend to be enriched in D as compared to their hosts, possibly due to higher rates of transpiration in the parasitic plants. The splitting of water in photosynthesis results in the lighter isotope being incorporated into organic matter. An even larger isotopic fractionation results during lipid synthesis and other processes involving the pyruvate dehydrogenase complex. Differences in metabolic pathway between species can be detected by D/H ratios. Hydrogen isotopic differences can be detected between CAM, C₄, and C₃ species. Within C₄ plants, the NADP-ME plants are isotopically distinguishable from NAD-ME and PEP-CK plants.     

Plant water uptake along a diversity gradient provides evidence for complementarity in hydrological niches

Biodiversity enhances a variety of ecosystem processes, and yet the underlying mechanisms through which these relationships occur remain a critical knowledge gap. Here, we used the natural abundance of stable isotopes to measure depth of water uptake in five common grassland species (Asclepias tuberosa, Lespedeza capitata, Liatris aspera, Schizachyrium scoparium and Sorghastrum nutans) growing across an experimental grassland diversity gradient. Using this approach, we addressed the following questions: 1) does the depth‐specific provenance of water uptake differ among species and/or do interspecific differences in water source manifest with increasing community diversity? 2) Does the isotopic niche space occupied by plants change with increasing diversity? 3) Is plasticity in water uptake depth across a diversity gradient associated with functional plant responses? We found that the depth of soil water used by plants was inherently different among species when grown in monocultures. All species used less shallow soil water and more intermediate‐depth soil water in mixed assemblages than in monocultures, resulting in similar interspecific differences in water source across the diversity gradient. However, plasticity in the locations of water used were positively associated with increases in plant growth in higher diversity treatments. These results indicate that plasticity in water‐use may contribute to positive biodiversity–productivity relationships commonly observed in temperate grasslands.     

Stable isotopes of carbon and nitrogen in soil ecological studies

The development of stable isotope techniques is one of the main methodological advances in ecology of the last decades of the 20th century. Many biogeochemical processes are accompanied by changes in the ratio between stable isotopes of carbon and nitrogen (¹²C/¹³C and ¹⁴N/¹⁵N), which allows different ecosystem components and different ecosystems to be distinguished by their isotopic composition. Analysis of isotopic composition makes it possible to trace matter and energy flows through biological systems and to evaluate the rate of many ecological processes. The main concepts and methods of stable isotope ecology and patterns of stable isotope fractionation during organic matter decomposition are considered with special emphasis on the fractionation of isotopes in food chains and the use of stable isotope studies of trophic relationships between soil animals in the field.     

Carbon isotopes in functional soil ecology

Soil is an integral part of terrestrial ecosystems. Many soil ecologists interested in soil ecosystem functioning rely, to some degree, on stable isotope methodologies. The study of the natural abundance of carbon isotopes, especially (13)C but also (14)C, in the environment and the use of stable carbon isotope tracers have proved very useful in investigating the soil carbon cycle and soil trophic relationships. Recent methodological and technical advances have greatly extended the possibilities for the application of stable carbon isotopes to terrestrial ecology and have vastly improved our knowledge of belowground ecosystem functioning and will continue to do so. A better understanding of soil processes is invaluable in predicting the future impacts of global environmental change on terrestrial ecosystems.     

长白山阔叶红松林演替序列植物-凋落物-土壤碳同位素特征

稳定碳同位素组成能精确指示生态系统碳循环过程,可以为深入研究森林演替进程对碳循环过程和固碳潜力的影响提供关键信息.利用稳定碳同位素技术对长白山阔叶红松林演替序列3种林分——中龄杨桦次生林、成熟杨桦次生林、阔叶红松林的叶片、树干、根系、凋落物和土壤δ¹³C值及碳、氮元素含量进行测定.结果表明: 各演替序列优势树种叶片δ¹³C从冠上到冠下均呈降低趋势;树干δ¹³C表现为树皮小于木质部;根系δ¹³C表现为细根小于粗根.阔叶红松林未分解凋落物δ¹³C小于半分解及全分解凋落物,次生林相反;土壤δ¹³C沿深度逐渐增加.总体上,δ¹³C值叶片<凋落物<根系<树干<土壤,说明植物各器官之间有明显的碳同位素分馏效应,且相同器官不同部位之间也存在差异;植物δ¹³C沿演替方向先减小后增加,土壤δ¹³C沿演替方向不断增加,且变化规律可以通过氮元素含量与碳同位素分馏效应的关系解释,说明长白山阔叶红松林演替过程优势树种和碳周转速率的变化影响了碳同位素分馏.     

Uncovering trophic positions and food resources of soil animals using bulk natural stable isotope composition

Despite the major importance of soil biota in nutrient and energy fluxes, interactions in soil food webs are poorly understood. Here we provide an overview of recent advances in uncovering the trophic structure of soil food webs using natural variations in stable isotope ratios. We discuss approaches of application, normalization and interpretation of stable isotope ratios along with methodological pitfalls. Analysis of published data from temperate forest ecosystems is used to outline emerging concepts and perspectives in soil food web research. In contrast to aboveground and aquatic food webs, trophic fractionation at the basal level of detrital food webs is large for carbon and small for nitrogen stable isotopes. Virtually all soil animals are enriched in ¹³C as compared to plant litter. This ‘detrital shift’ likely reflects preferential uptake of ¹³C‐enriched microbial biomass and underlines the importance of microorganisms, in contrast to dead plant material, as a major food resource for the soil animal community. Soil organic matter is enriched in ¹⁵N and ¹³C relative to leaf litter. Decomposers inhabiting mineral soil layers therefore might be enriched in ¹⁵N resulting in overlap in isotope ratios between soil‐dwelling detritivores and litter‐dwelling predators. By contrast, ¹³C content varies little between detritivores in upper litter and in mineral soil, suggesting that they rely on similar basal resources, i.e. little decomposed organic matter. Comparing vertical isotope gradients in animals and in basal resources can be a valuable tool to assess trophic interactions and dynamics of organic matter in soil. As indicated by stable isotope composition, direct feeding on living plant material as well as on mycorrhizal fungi is likely rare among soil invertebrates. Plant carbon is taken up predominantly by saprotrophic microorganisms and channelled to higher trophic levels of the soil food web. However, feeding on photoautotrophic microorganisms and non‐vascular plants may play an important role in fuelling soil food webs. The trophic niche of most high‐rank animal taxa spans at least two trophic levels, implying the use of a wide range of resources. Therefore, to identify trophic species and links in food webs, low‐rank taxonomic identification is required. Despite overlap in feeding strategies, stable isotope composition of the high‐rank taxonomic groups reflects differences in trophic level and in the use of basal resources. Different taxonomic groups of predators and decomposers are likely linked to different pools of organic matter in soil, suggesting different functional roles and indicating that trophic niches in soil animal communities are phylogenetically structured. During last two decades studies using stable isotope analysis have elucidated the trophic structure of soil communities, clarified basal food resources of the soil food web and revealed links between above‐ and belowground ecosystem compartments. Extending the use of stable isotope analysis to a wider range of soil‐dwelling organisms, including microfauna, and a larger array of ecosystems provides the perspective of a comprehensive understanding of the structure and functioning of soil food webs.     

高寒草甸不同演替阶段植物叶片功能性状研究

以甘南高寒草甸演替过程中5个阶段的典型群落为研究对象,对围封样地内不同演替阶段群落叶片功能性状进行比较,分析了围封地内各演替阶段群落水平上主要物种叶性状与环境因子的关系。结果显示:(1)此围封地形成了一个从草本到灌木的演替过程。随着演替的进行,Margalef指数显著增加,Shannon-Wiener指数呈先增加后降低的趋势。(2)从演替前期到演替后期,土壤有机碳(SOC)、土壤全氮(STN)、土壤含水量(SWC)逐渐升高,光照度(LI)、土壤温度(ST)逐渐降低,土壤全磷(STP)呈先降低后增加趋势;叶片有机碳(LCC)、全氮(LNC)、含水量(LWC)逐渐升高;比叶面积(SLA)、磷利用效率(PUPE)、稳定碳同位素(δ13C)逐渐下降,叶片全磷(LPC)先降低后升高,而氮利用效率(PUNE)先升高后降低。(3)RDA冗余分析表明,在此围封样地内,演替前期植物群落叶性状主要受到LI和ST的限制作用。而在演替的中后期SWC[WTBZ]是主要影响因子。此研究有助于我们认识高寒草甸生态系统的退化过程所导致的生态环境问题,进而寻求更好的草地恢复和重建方法。