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。     

箱式通量观测技术和方法的理论假设及其应用进展

碳(CO₂、CH₄)、氮(N₂O)和水汽(H₂O)等温室气体的交换通量是生态系统物质循环的核心, 是地圈-生物圈-大气圈相互作用的纽带。稳定同位素光谱和质谱技术和方法的进步使碳稳定同位素比值(δ ¹³C)和氧稳定同位素比值(δ ¹⁸O)(CO₂)、δ ¹³C (CH₄)、氮稳定同位素比值(δ ¹⁵N)和δ ¹⁸O (N₂O)、氢稳定同位素比值(δD)和δ ¹⁸O (H₂O)的观测成为可能, 与箱式通量观测技术和方法结合可以实现土壤、植物乃至生态系统尺度温室气体及其同位素通量观测研究。该综述以CO₂及其δ ¹³C通量的箱式观测技术和方法为例, 概述了箱式通量观测系统的基本原理及分类, 阐述了系统设计的理论要求和假设, 综述了从野外到室内土壤、植物叶-茎-根以及生态系统尺度箱式通量观测研究的应用进展及问题, 展望了气体分析精度和准确度、观测数据精度和准确度以及观测数据的代表性评价在箱式通量观测研究中的重要性。     

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。     

碳同位素示踪技术及其在陆地生态系统碳循环研究中的应用与展望

碳同位素示踪技术具有高度的专一性和灵敏度, 经过几十年的发展, 形成了一系列成熟的标记方法, 在陆地生态系统碳循环过程的研究中已得到广泛应用。目前, 自然丰度法、与¹³C贫化示踪技术结合的自由空气中气体浓度增加(FACE)实验、脉冲与连续标记法以及碳同位素高丰度底物富集标记法是研究陆地生态系统碳循环过程常用的碳同位素示踪方法; 通过将长期定位实验和室内模拟实验结合, 量化光合碳在植物-土壤系统的传输与分配特征, 明确植物光合碳对土壤有机质的来源、稳定化过程的影响及其微生物驱动机制; 阐明土壤碳动态变化(迁移与转化)和新碳与老碳对土壤碳库储量的相对贡献, 评估有机碳输入、转化与稳定的生物与非生物微观界面过程机制。然而, 生态系统碳循环受气候、植被、人为活动等多因素影响, 碳同位素技术需要结合质谱、光谱技术实现原位示踪, 结合分子生物学技术阐明其微生物驱动机制, 从而构建灵敏、准确、多尺度、多方位的同位素示踪技术体系。因此, 该文以稳定碳同位素为主, 综述了碳同位素示踪技术的原理、分析方法和在陆地生态系统碳循环过程中的应用进展, 归纳总结了碳同位素示踪技术结合原位检测技术和分子生物学技术的研究进展和应用前景, 并对碳同位素示踪技术存在的问题进行了分析和展望。     

Hydrogen isotopic compositions of n-alkanes from terrestrial plants correlate with their ecological life forms

Stable hydrogen isotopic compositions (δD) of compound-specific biomarkers, such as n-alkanes from plant leaf waxes, can be used as a proxy for paleoclimatic change. However, the relationship between hydrogen isotopes of plant leaf wax and plant ecological life forms is not well understood. Here, we report the δD of n-alkanes from 34 modern terrestrial plants, including twenty-one C₃ plants and thirteen C₄ plants from northwestern China, determined using gas chromatography/thermal conversion/isotope ratio mass spectrometry. Our data show that the stable hydrogen isotopes are poorly correlated with the plant photosynthetic pathway (C₃ vs. C₄) and that they do not give clear regional precipitation signals. Together with a comparative analysis of published δD values from plant leaf waxes in other regions, we believe that the stable hydrogen isotope of plant leaf waxes is more closely related to ecological life forms of these terrestrial plants (i.e. tree, shrub, and grass). In general, the grasses have more negative δD values than the co-occurring trees and shrubs. Our findings suggest that the δD values of sedimentary leaf waxes from higher plants may record changes of a plant ecosystem under the influence of environmental alteration and imply that reconstruction of the paleoclimate using δD values from plant n-alkanes should be based upon specific plant taxa, and comparison should be made among plants with similar ecological life forms.     

Shrub encroachment effect on the evapotranspiration and its component—A numerical simulation study of a shrub encroachment grassland in Nei Mongol,China

Aims Shrub encroachment is a common global change phenomenon occurring in arid and semi-arid regions. Due to the difficulty of partitioning evapotranspiration into shrub plants, grass plants and soil in the field, there are few studies focusing on shrub encroachment effect on the evapotranspiration and its component in China. This study aims to illustrate shrub encroachment effect on evapotranspiration by the numerical modeling method. Methods A two-source model was applied and calibrated with the measured evapotranspiration (ET) by the Bowen ratio system to simulate evapotranspiration and its component in a shrub encroachment grassland in Nei Mongol, China. Based on the calibrated model and previous shrub encroachment investigation, we set three scenarios of shrub encroachment characterized by relative shrub coverage of 5%, 15% and 30%, respectively, and quantified their effects caused by shrub encroachment through localized and calibrated two-source model.Important findings The two-source model can well reconstruct the evapotranspiration characteristics of a shrub encroachment grassland. Sensitivity analysis of the model shows that errors for the input variables and parameters have small influence on the result of partitioning evapotranspiration. The result shows that shrub encroachment has relatively small influence on the total amount of ET, but it has clear influence on the proportion of the components of evapotranspiration (E/ET). With shrub coverage increasing from 5% to 15% and then 30%, the evapotranspiration decreased from 182.97 to 180.38 and 176.72 W·m^-2, decreasing amplitude values of 0.34% and 0.44%, respectively. On average, E/ET rises from 52.9% to 53.9% and 55.5%, increasing amplitude values to 2.04% and 3.25%. Data analysis indicates that shrub encroachment results in smaller soil moisture changes, but clear changes of ecosystem structure (decreasing ecosystem leaf area index while increasing vegetation height) which lead to the decrease of transpiration fraction through decreasing canopy conductance. The research highlights that, with the shrub encroachment, more water will be consumed as soil evaporation which is often regarded as invalid part of evapotranspiration and thus resulting in the decrease of water use efficiency.     

基于多种同位素模型的侧柏林生态系统蒸散组分定量拆分

为了全面认识森林生态系统蒸散各组分及其对蒸散的贡献率在日尺度上的变化规律,本研究利用同位素稳态和非稳态假设理论结合水同位素分析仪系统,对生长季侧柏林生态系统蒸散各组分进行了定量拆分和比较。结果表明: 4个测定日(2016年8月5、8、10、11日)不同来源水体的¹⁸O都呈现表层土壤水氧同位素组成(δ_S)＞枝条水氧同位素组成(δ_X)＞大气水汽氧同位素组成(δ_V),说明三者可能因同位素分馏效应表现出明显的差异。土壤蒸发水汽氧同位素组成(δ_E)在日尺度上为-26.89‰～-59.68‰,整体上呈现出先上升后下降的变化趋势;森林生态系统蒸散水汽氧同位素组成(δ_ET)为-15.99‰～-10.04‰,稳态(ISS)下植物蒸腾水汽氧同位素组成(δ_T-ISS)为-12.10‰～-9.51‰,而非稳态(NSS)下植物蒸腾水汽氧同位素组成(δ_T-NSS)为-13.02‰～-7.23‰,在日时间尺度上δ_ET与δ_T-NSS全天的变化趋势一致,在11:00—17:00 δ_ET、δ_T-ISS与δ_T-NSS三者的变化趋势近似一致。总体上,植物蒸腾量对蒸散量的贡献率表现为F_T-ISS 79.1%～98.7%,而F_T-NSS 88.7%～93.7%。这表明研究区土壤蒸发耗水远小于植被蒸腾耗水,植被蒸腾在林地蒸散中起主导作用。     

黑河中游春玉米叶片水δD和δ18O的富集过程和影响因素

植物水的稳定同位素分馏过程是水在土壤-植物-大气连续体中循环的重要环节。以往研究由于叶片水¹⁸O同位素比值(δ¹⁸O _l,b)和氘(D)同位素比值(δD_l,b)(合称δ_l,b)实测数量少只能作为模型验证数据, 导致δ_l,b富集机制研究多集中于模型研究, 缺乏基于野外试验条件的δ_l,b富集的控制机制研究。叶片水δD_l,b和δ¹⁸O _l,b的富集程度(ΔD_l,b和Δ¹⁸O _l,b, 合称Δ_l,b)通常表示为δ_l,b与茎秆水D同位素比值(δD_x)和¹⁸O同位素比值(δ¹⁸O_x) (合称δ_x)之差, 即Δ_l,b = δ_l,b - δ_x。该研究以黑河中游沙漠绿洲春玉米(Zea mays)生态系统为研究对象, 重点采集和分析了季节和日尺度δ_l,b和δ_x数据, 配套开展了大气水汽δ¹⁸O和δD (合称δ_v)等辅助变量的原位连续观测, 探讨了季节和日尺度上的δ_l,b富集特征及其影响因素。结果表明: 叶片水δ_l,b和Δ_l,b的季节变化趋势不明显, 而受蒸腾作用影响表现出白天富集夜间贫化的单峰日变化特征。对于D来说, 无论季节尺度上还是日尺度上, 大气水汽δ_v和相对湿度是δD_l,b和ΔD_l,b的主要环境控制因素; 而对于¹⁸O来说, 无论季节尺度上还是日尺度上, 相对湿度是δ¹⁸O _l,b和Δ¹⁸O _l,b的主要环境控制因素。由于D和¹⁸O在热力学平衡分馏上有约8倍差异, 直接分析叶片水ΔD_l,b和Δ¹⁸O_l,b与影响因素的差异性, 有助于理解叶片水δD和δ¹⁸O富集过程以及对模型发展有一定的指导意义。     

基于稳定同位素的SPAC水碳拆分及耦合研究进展

土壤-植被-大气连续体(SPAC)是陆地水文学、生态学和全球变化领域的重要研究对象,其水碳循环过程及耦合机制是前沿性问题.稳定同位素技术示踪、整合和指示的特征有助于评估分析生态系统固碳和耗水情况.本文在简述稳定同位素应用原理和技术的基础上,重点阐释了基于稳定同位素光学技术的SPAC系统水碳交换研究进展,包括:在净碳通量中拆分光合与呼吸量,在蒸散通量中拆分蒸腾与蒸发量,以及在系统尺度上的水碳耦合研究.新兴的技术和方法实现了生态系统尺度上长期高频的同位素观测,但在测量精准度、生态系统呼吸拆分、非稳态模型适应性、尺度转换和水碳耦合机制等方面存在挑战.本文探讨了现有主要研究成果、局限性以及未来研究展望,以期对稳定同位素生态学领域的新研究和技术发展有所帮助.     

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。     

Stable isotopes in leaf water of terrestrial plants

Leaf water contains naturally occurring stable isotopes of oxygen and hydrogen in abundances that vary spatially and temporally. When sufficiently understood, these can be harnessed for a wide range of applications. Here, we review the current state of knowledge of stable isotope enrichment of leaf water, and its relevance for isotopic signals incorporated into plant organic matter and atmospheric gases. Models describing evaporative enrichment of leaf water have become increasingly complex over time, reflecting enhanced spatial and temporal resolution. We recommend that practitioners choose a model with a level of complexity suited to their application, and provide guidance. At the same time, there exists some lingering uncertainty about the biophysical processes relevant to patterns of isotopic enrichment in leaf water. An important goal for future research is to link observed variations in isotopic composition to specific anatomical and physiological features of leaves that reflect differences in hydraulic design. New measurement techniques are developing rapidly, enabling determinations of both transpired and leaf water δ¹⁸O and δ²H to be made more easily and at higher temporal resolution than previously possible. We expect these technological advances to spur new developments in our understanding of patterns of stable isotope fractionation in leaf water.     

Ecological thresholds: Concept,Methods and research outlooks

The concept of ecological thresholds was raised in the 1970s. However, it was subsequently given different definitions and interpretations depending on research fields or disciplines. For most scientists, ecological thresholds refer to the points or zones that link abrupt changes between alternative stable states of an ecosystem. The measurement and quantification of ecological thresholds have great theoretical and practical significance in ecological research for clarifying the structure and function of ecosystems, for planning sustainable development modes, and for delimiting ecological red lines in managing the ecosystems of a region. By reviewing the existing concepts and classifications of ecological thresholds, we propose a new concept and definition at two different levels: the ecological threshold points, i.e. the turning points of quantitative changes to qualitative changes, which can be considered as ecological red lines; the ecological threshold zones, i.e. the regime shifts of the quantitative changes among different stable states, which can be considered as the yellow and/or orange warning boundaries of the gradual ecological changes. The yellow thresholds mean that an ecosystem can return to a stable state by its self-adjustment, the orange thresholds indicate that the ecosystem will stay in the equilibrium state after interference factors being removed, whereas the red thresholds, as the critical threshold points, indicate that the ecosystem will undergo irreversible degradation or even collapse beyond those points. We also summarizes two types of popular Methods in determining ecological thresholds: statistical analysis and modeling based on data of field observations. The applications of ecological thresholds in ecosystem service, biodiversity conservation and ecosystem management research are also reviewed. Future research on ecological thresholds should focus on the following aspects: (1) methodological development for measurement and quantification of ecological thresholds; (2) emphasizing the scaling effect of ecological thresholds and establishment of national-scale observation system and network; and (3) implementation of ecological thresholds as early warning tools in ecosystem management and delimiting ecological red lines.     

植物水分来源稳定氢氧同位素偏移研究进展

稳定氢氧同位素技术能有效计算植物根系水分吸收量,确定植物水分来源贡献,评估植物水分利用策略,是生态水文学探究大气-植被-土壤系统水分传输过程机制的有效工具。然而土壤与木质部水稳定氢氧同位素比值(δ²H和δ¹⁸O)偏移造成植物水分来源贡献率计算偏差,引起氢氧同位素结果差异的原因尚不明晰。该文首先简要介绍氢氧稳定同位素比值偏移现象,其次沿水分在土壤-植物-大气连续体中的传输路径构建梳理框架,系统阐述了3个界面(植物-大气界面、土壤-大气界面和根系-土壤界面)与2个空间(植物体和土壤层)中引起δ²H与δ¹⁸O偏移的自然效应,同时概述了土壤与木质部样品提取与测定技术中引起δ²H与δ¹⁸O偏差的人为效应。最后,根据现有研究进展提出主要问题,从获取同位素时空数据,微尺度同位素偏移原因,提取与测定技术的优化三方面指出未来的发展方向。     

碳同位素技术在森林生态系统碳循环研究中的应用

碳同位素技术对碳素在生态系统中的迁移动态具有很好的示踪作用,在生态学各领域研究中应用广泛。土壤、大气、植物是森林生态系统的重要碳库,植物是大气和土壤交换碳元素的重要介质。本文简要总结了碳同位素技术在研究碳元素在植物体内以及植物、土壤、大气碳库之间的迁移规律和生态学过程中的应用,展望了该技术在森林界面学中的应用前景。     

Dominant species and ecosystem gas exchange in temperate grassland under different land use patterns

在内蒙古温带草原围封、放牧和割草3种处理下的样地内, 对生态系统尺度和大针茅(Stipa grandis)、冷蒿(Artemisia frigida)、羊草(Leymus chinensis) 3种优势种植物叶片尺度上的气体交换和水分关系进行了测定, 对比研究了植物碳水对环境的响应。结果表明, 在优势种单株尺度和生态系统尺度上, 大气-植被CO₂交换因草地利用方式的不同而具有不同的表现。在生态系统层面, 放牧样地的群落净CO₂气体交换量和总初级生产力都与围封样地和割草样地有差异, 群落总初级生产力受生态系统呼吸的影响。在放牧处理下, 群落净CO₂气体交换量日变化表现为生态系统对碳的吸收, 而围封和割草则以碳释放为主。单叶光合速率出现负值并随时间推移而恢复的现象, 应是植物对干旱高温、高光照的特殊反应。生态系统水分利用效率没有明显不同, 但各样地的蒸散能力有趋势上的变化; 对于同种植物, 放牧样地植物单叶水分利用效率的日变化波动幅度最大, 围封样地最小。     

利用稳定氢氧同位素定量区分白刺水分来源的方法比较

水是影响植物分布的重要生态因子之一,对植物水源的研究有助于在全球变化背景下了解植物的时空分布格局.根据同位素质量守恒,利用稳定氢氧同位素可以确定植物水分来源,相关的方法也不断改进.利用三源线性混合模型、多源线性混合模型、吸水深度模型以及动态模型分别对格尔木白刺(Nitraria Tangutorum)的水分来源进行了对比研究,发现格尔木白刺主要吸收利用50-100 cm处的土壤水及地下水.在研究方法上,各模型都有自己的应用范围和局限:三源线性混合模型一般只能在植物吸收的水分来源不超过3个的情况下运行;多源线性混合模型弥补了三源线性混合模型的不足,可以同时比较多种来源水各自对白刺的贡献率及贡献范围;吸水深度模型弥补了混合模型中不能计算白刺对土壤水的平均吸水深度的缺陷;动态模型则会为未来降水格局变化对植物的时空分布的影响研究起很大作用.针对不同的适用范围,模型的选择及综合应用会更广泛.但是,该技术还存在一些不足,需要结合测定土水势,富氘水的示踪等方法来弥补.     

Effects of environmental parameters, leaf physiological properties and leaf water relations on leaf water delta18O enrichment in different Eucalyptus species

Stable oxygen isotope ratios (delta18O) have become a valuable tool in the plant and ecosystem sciences. The interpretation of delta18O values in plant material is, however, still complicated owing to the complex interactions among factors that influence leaf water enrichment. This study investigated the interplay among environmental parameters, leaf physiological properties and leaf water relations as drivers of the isotopic enrichment of leaf water across 17 Eucalyptus species growing in a common garden. We observed large differences in maximum daily leaf water delta18O across the 17 species. By fitting different leaf water models to these empirical data, we determined that differences in leaf water delta18O across species are largely explained by variation in the Péclet effect across species. Our analyses also revealed that species-specific differences in transpiration do not explain the observed differences in delta18O while the unconstrained fitting parameter 'effective path length' (L) was highly correlated with delta18O. None of the leaf morphological or leaf water related parameters we quantified in this study correlated with the L values we determined even though L was typically interpreted as a leaf morphological/anatomical property. A sensitivity analysis supported the importance of L for explaining the variability in leaf water delta18O across different species. Our investigation highlighted the importance of future studies to quantify the leaf properties that influence L. Obtaining such information will significantly improve our understanding of what ultimately determines the delta18O values of leaf water across different plant species.     

Leaf water 18O and 2H enrichment along vertical canopy profiles in a broadleaved and a conifer forest tree

Distinguishing meteorological and plant‐mediated drivers of leaf water isotopic enrichment is prerequisite for ecological interpretations of stable hydrogen and oxygen isotopes in plant tissue. We measured input and leaf water δ²H and δ¹⁸O as well as micrometeorological and leaf morpho‐physiological variables along a vertical gradient in a mature angiosperm (European beech) and gymnosperm (Douglas fir) tree. We used these variables and different enrichment models to quantify the influence of Péclet and non‐steady state effects and of the biophysical drivers on leaf water enrichment. The two‐pool model accurately described the diurnal variation of leaf water enrichment. The estimated unenriched water fraction was linked to leaf dry matter content across the canopy heights. Non‐steady state effects and reduced stomatal conductance caused a higher enrichment of Douglas fir compared to beech leaf water. A dynamic effect analyses revealed that the light‐induced vertical gradients of stomatal conductance and leaf temperature outbalanced each other in their effects on evaporative enrichment. We conclude that neither vertical canopy gradients nor the Péclet effect is important for estimates and interpretation of isotopic leaf water enrichment in hypostomatous trees. Contrarily, species‐specific non‐steady state effects and leaf temperatures as well as the water vapour isotope composition need careful consideration.     

Research progress on water uptake through foliage

Leaf is one of the most sensitive organs that response to atmospheric environment changes. In arid region, especially under water stress conditions, water from foliar uptake is one of the main water subsidy sources, which plays an important role in plant survival, growth and ecosystem sustainable development. Foliar water uptake attracts more attention in physiological and ecological research all over the world. This essay summarized the main measurement methods of foliar water uptake at home and abroad in detail: vessel control method, stable isotope technology and heat ratio method. And analyzed the significance of foliar water uptake from physiological and ecological aspects; and discussed the influence factors of foliar water uptake from water condition perspective, water absorption and leaf structure aspects. The following aspects should be paid more attention in future researches: (1) measure the amount of foliar water uptake by using stable isotope technology and heat ratio method; (2) the relationship among foliar uptake of fog, dew, rain and water vapor in arid region; (3) the foliar uptake amount and rate should be simulated by models; (4) the mechanisms of foliar water uptake should be researched; (5) study foliar water uptake of different plants at ecosystem scale.     

植物水分利用策略研究进展

水分是影响植物生长发育的重要因子之一。地球上大多数生态系统中的植物都会经历一个降水相对稀少的干旱季节,植物在不同的季节与不同的生态系统中究竟如何利用水分,利用哪些水分去获得生存,成为一个人们关注的问题。在过去的20年,稳定同位素技术在植物生态学中的应用得到了稳定长足的发展。因为陆地植物(少数排盐种类除外)在水分吸收过程中不发生同位素分馏,因此可以利用δD与δ18O数据进行水分获取方式的研究。对植物木质部水分以及其潜在水源的稳定同位素进行分析,并参考土壤水势、叶片水势、土壤含水量等数据,同时运用二元或三元混合模型,可以定量确定植物的水分利用来源。大量的研究表明,不同功能型、生长阶段、季节的植物以及不同物种往往具有不同的水分利用策略。