基于稳定同位素的干旱半干旱区SPAC水分运移过程研究进展

土壤-植物-大气连续体(SPAC)是生态水文学的重点研究对象,其水分运移过程对于干旱半干旱区生态植被建设和水资源综合管理具有重要意义。氢氧稳定同位素较高的灵敏性和准确度有助于揭示这一过程。介绍了氢氧稳定同位素在土壤-大气界面、土壤-地下水界面、土壤-植物界面和植物-大气界面水分补给传输过程中的应用,包括土壤水分来源和蒸发;水分补给入渗机制和滞留时间;植物水分来源和水力再分配;蒸散发分割和叶片吸水的相关研究,同时明确了氢氧稳定同位素技术在应用过程中存在的一些不确定性以及未来亟需加强的方面,以期为利用稳定同位素技术对生态水文过程的研究提供参考依据。     

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。     

亚热带地区常绿阔叶林SPAC系统水分的氢氧稳定同位素特征

森林的土壤-植物-大气连续体(SPAC)是陆地重要的水循环连续界面过程。本研究通过分析亚热带常绿阔叶林的降水、大气水汽、土壤水、叶片水的同位素组成,探讨森林SPAC系统水分的氢氧同位素组成特征以及植物蒸腾与叶片性状和环境因子的关系。结果表明: 研究区大气降水、土壤水、竹柏枝条水、竹柏叶片水和大气水汽的δD-δ¹⁸O线性回归方程分别为: δD_P=7.97δ¹⁸O_P+12.68(R²=0.97)、δD_S=4.29δ¹⁸O_S-18.62(R²=0.81)、δD_B=3.31δ¹⁸O_B-29.73(R²=0.49)、δD_L=1.49δ¹⁸O_L-10.09(R²=0.81)、δD_V=3.89δ¹⁸O_V-51.29(R²=0.46)。在降水→土壤水→植物水的界面水输送过程中,氢氧同位素逐渐富集,而从土壤蒸发和从植物蒸腾的水汽同位素贫化。在降水和蒸发作用的影响下,土壤水同位素随深度增加有贫化的趋势,而且整体上旱季土壤水同位素比雨季富集。观测期间,枝条水同位素比土壤水略微富集,说明水分在植物体内运输过程中存在受到蒸腾富集作用的可能性。旱季,乔木的枝条水同位素比灌木贫化,说明根系分布更深的乔木植物更倾向于利用深层土壤水。由于在叶片性状、蒸腾速率以及对环境因子的响应程度等方面存在差异,不同植物的叶片水同位素组成随叶龄增长的变化特征有所不同。雨季的环境条件更有利于叶片蒸腾,使雨季的叶片水同位素比旱季富集。叶片水同位素组成与植物叶片含水量呈正相关关系,与相对湿度呈负相关关系,综合反映了植物应对环境变化的水分调控功能。     

基于稳定氧同位素确定植物水分来源不同方法的比较

利用稳定同位素技术确定植物水分来源,对提高生态水文过程的认识和对干旱半干旱区的生态管理至关重要。目前基于稳定同位素技术确定植物水分来源的方法众多,但不同方法之间对比的研究较少。本研究基于原位样品采集,室内实验测试,利用直接对比法、多元线性混合模型(IsoSource)、贝叶斯混合模型(MixSIR、MixSIAR)和吸水深度模型分析植物水分来源,并对比各方法的优缺点。结果表明:相对于多元线性混合模型(IsoSource)而言,贝叶斯混合模型(MixSIR、MixSIAR)具有更好的水源区分性能,但对数据要求较高,且植物木质部水和潜在水源同位素组成的标准差越小,模型运行结果的可信度更高。本研究中贝叶斯混合模型(MixSIR)为最优解。在利用稳定氢氧同位素技术确定植物水分来源时,可先通过直接对比法定性判断植物可能利用的潜在水源,然后再用多元线性混合模型(IsoSource)、贝叶斯混合模型(MixSIR、MixSIAR)计算出各潜在水源对植物的贡献率和贡献范围,必要时可评估模型性能,选择出最优模型,定量分析植物的水分来源。若植物主要吸收利用不同土层深度的土壤水,可结合吸水深度模型计算出植物吸收土壤水的平均深度。本研究为干旱半干旱地区利用同位素技术确定植物水分来源方法的选择提供了理论依据。     

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 decomposition and storage conditions on the δ13C and δ15N isotope values of killer whale (Orcinus orca) skin and blubber tissues

Several different factors in the collection and preservation of whale skin and blubber samples were examined to determine their effect on the results obtained by stable nitrogen and carbon isotope (δ¹⁵N and δ¹³C) analysis. Samples of wet killer whale skin retained their original stable isotope values for up to 14 d at 4°C or lower. However, decomposition significantly changed the δ¹⁵N value within 3 d at 20°C. Storage at ?20°C was as effective as ?80°C for the preservation of skin and blubber samples for stable isotope analysis for at least a year. By contrast, once a skin sample had been freeze‐dried and lipid extracted, the stable isotope values did not change significantly when it was stored dry at room temperature for at least 12 mo. Preservation of whale skin samples for a month in DMSO‐salt solution, frozen or at room temperature, did not significantly change the δ¹⁵N and δ¹³C values of lipid extracted tissues, although the slight changes seen could influence results of a study if only small changes are expected.     

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。     

不同降水条件下两种荒漠植物的水分利用策略

自然降水是干旱、半干旱地区荒漠植物重要的水分来源。为了说明自然降水量的变化对干旱、半干旱地区荒漠植物水分利用策略的影响, 研究了两种常见荒漠植物油蒿(Artemisia ordosica)和白刺(Nitraria tangutorum)在3个不同自然降水地区(内蒙古的杭锦旗和磴口县及甘肃的民勤县)的水分来源、水分利用效率及植物的抗逆能力的变化。测定了不同地区的植物茎水、各潜在水源(降水、地下水和土壤水)的δD和δ¹⁸O值, 并利用IsoSource模型分析了这两种植物在不同地区对这些潜在水源的选择性利用情况; 同时测定了叶片的δ¹³C和游离脯氨酸浓度。结果表明: 在年降水量最高的杭锦旗, 这两种植物对浅层土壤水的利用比例最高, 其中油蒿主要利用0-50 cm土层中的水源; 在年降水量相对较低的磴口和民勤, 植物利用的主要水源为深层土壤水和地下水。随着年降水量的增加, 这两种植物的水分利用效率逐渐降低。白刺的脯氨酸浓度大于油蒿, 与水分利用效率无关, 但油蒿的水分利用效率和脯氨酸浓度成正比。研究表明, 荒漠植物能通过改变其水分利用策略和其他生理特性适应自然降水量的变化, 但不同植物种采用的策略可能有所不同。     

准噶尔盆地东南缘多枝柽柳、白刺和红砂水分来源的异同

荒漠生态系统中, 水是植物生长最主要的限制因子。为了比较同一生境下不同荒漠植物的水分来源特征, 选取了同一生境下的多枝柽柳(Tamarix ramosissima)、白刺(Nitraria sibirica)和红砂(Reaumuria soongorica), 测定了这3种植物茎水和各潜在水源(降水、土壤水和地下水)的氢、氧稳定同位素比率(δD和δ¹⁸O)值, 并利用IsoSource软件计算了3种植物对潜在水源的利用比例。结果表明: 红砂和白刺的茎水δD和δ¹⁸O值及其水分来源有明显的季节波动特征。其中, 红砂为浅根系植物, 春季(3-5月)以表层土壤水为主要水源, 夏秋季节(6-10月)表层土壤含水量显著降低, 其主要的水分来源逐渐偏向于较深层的土壤水; 白刺的根系分布范围介于红砂和多枝柽柳之间, 在春季能够较多地利用表层土壤水, 而到了夏秋季节, 所利用的水分更多地来源于深层土壤水或地下水; 多枝柽柳为深根系植物, 其90%以上的水分来源于深层土壤水和地下水, 而且茎水δD和δ¹⁸O值及其水分来源没有季节波动特征。3种植物水分来源特征的差异与其水分利用策略密切相关, 同时, 也说明荒漠灌木可以通过自身调节向着最优(最有利)表现型发展, 从而最大限度地获取水分。     

石灰岩地区连片出露石丛生境植物水分来源的季节性差异

以桂西北石灰岩地区连片出露石丛生境次生林群落为研究对象, 运用稳定性氢氧同位素技术结合IsoSource模型, 分析了5种典型植物(半落叶乔木粉苹婆(Sterculia euosma)、落叶乔木菜豆树(Radermachera sinica)、常绿灌木鹅掌柴(Schefflera octophylla)以及落叶灌木红背山麻杆(Alchornea trewioides)和紫弹树(Celtis biondii))水分来源的季节性差异。结果表明: 雨季, 除鹅掌柴同时利用部分前期雨水外, 其余4种植物均主要利用最近15天内的雨水(利用比例的平均值之和均超过80%)。旱季, 5种植物均主要利用最近一次雨水与前期雨水(一个月前)的混合(利用比例的平均值之和均超过80%), 其中乔木和常绿灌木对前期雨水的利用比例更高(利用比例的平均值均超过50%)。植被良好的石灰岩连片出露石丛生境中, 植物主要以不同时期的雨水为主要水源, 这可能与石灰岩发育的裂隙及其储水能力有关。储存在裂隙中的雨水通过植物蒸腾的方式返回大气, 这一良好的水文循环过程应得到充分的保护。     

西双版纳绞杀植物斜叶榕的水分利用策略

以斜叶榕为研究对象,通过测定其不同生长阶段木质部与各潜在水源的稳定氢、氧同位素组成,以及土壤水分含量、土壤水势、叶片水势等参数,揭示西双版纳地区不同生长阶段的绞杀榕（斜叶榕）在不同季节的水分利用策略.结果表明：浅层土壤（10～50 cm）的水势在干热季与雾凉季变化较大,较深土壤（51～120 cm）的水势在各季节无明显变化;雾凉季与干热季的土壤含水量之间无显著差异（P=0.64）;植物黎明前叶片水势与中午叶片水势随不同生长阶段而异;根据木质部水与各潜在水源的稳定氧同位素以及植物水势等其他参数判定,浅层土壤水是斜叶榕全年最主要的水分来源,不同生长阶段的斜叶榕在不同季节采取了不同的水分利用策略来应对环境的变化.     

Nitrogen and carbon isotope variability in the green‐algal lichen Xanthoria parietina and their implications on mycobiont–photobiont interactions

Stable isotope patterns in lichens are known to vary largely, but effects of substrate on carbon and nitrogen stable isotope signatures of lichens were previously not investigated systematically. N and C contents and stable isotope (δ¹⁵N, δ¹³C) patterns have been measured in 92 lichen specimens of Xanthoria parietina from southern Bavaria growing on different substrates (bark and stone). Photobiont and mycobiont were isolated from selected populations and isotopically analyzed. Molecular investigations of the internal transcribed spacer of the nuclear ribosomal DNA (ITS nrDNA) region have been conducted on a subset of the specimens of X. parietina. Phylogenetic analysis showed no correlation between the symbionts X. parietina and Trebouxia decolorans and the substrate, isotope composition, or geographic origin. Instead specimens grown on organic substrate significantly differ in isotope values from those on minerogenic substrate. This study documents that the lichens growing on bark use additional or different N sources than the lichens growing on stone. δ¹⁵N variation of X. parietina apparently is controlled predominantly by the mass fraction of the mycobiont and its nitrogen isotope composition. In contrast with mycobionts, photobionts of X. parietina are much more ¹⁵N‐depleted and show less isotopic variability than mycobionts, probably indicating a mycobiont‐independent nitrogen acquisition by uptake of atmospheric ammonia.     

The nutritional significance of a winter‐flowering succulent for opportunistic avian nectarivores

The winter‐flowering succulent Aloe marlothii provides nectar for many opportunistic avian nectarivores in southern African savannas. We assessed the importance of A. marlothii nectar sugar for opportunistic nectarivores by analysing temporal changes in stable carbon isotope ratios (δ¹³C) in the tissues of birds in Suikerbosrand Nature Reserve, South Africa. The blood of the 11 most common non‐granivorous opportunistic nectarivores at our site was enriched in ¹³C by 3.4 ± 1.5‰ during the flowering period of A. marlothii, reflecting the enriched crassulacean acid metabolism (CAM) isotopic signature of nectar (?12.6 ± 0.5‰). This relatively small contribution of A. marlothii nectar to assimilated carbon in whole blood contrasted with that of exhaled CO₂ in African Red‐eyed Bulbuls Pycnonotus nigricans and Cape White‐eyes Zosterops capensis. In both these species, the δ¹³C of breath samples was significantly enriched compared with blood and feathers, and closely resembled that of the nectar, revealing combustion of ingested nectar rather than assimilation. Although our analysis was complicated by the presence of C₄ grasses, whose δ¹³C values are similar to those of CAM photosynthesizers, when considered with previously published feeding observations our data reveal that opportunistic nectarivores feeding on A. marlothii nectar obtain a relatively small fraction of their assimilated carbon, but most of their metabolized carbon, from this seasonally available carbohydrate food resource. Because the δ¹³C values of insects associated with C₃ plants also became enriched during the flowering season, some insect‐eating opportunistic nectarivores may have assimilated A. marlothii carbon indirectly from insects. This study highlights the importance of understanding isotopic routing when assessing the nutritional significance of specific dietary items to consumer communities.     

Dual‐tracer‐based isotope turnover rates in a highly invasive mysid Limnomysis benedeni from Lake Constance

Understanding the ecological patterns of invasive species and their habitats require an understanding of the species’ foraging ecology. Stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope values provide useful information into the study of animal ecology and evolution, since the isotope ratios of consumers reflect consumer's dietary patterns. Nevertheless, the lack of species‐ and element‐specific laboratory‐derived turnover rates could limit their application. Using a laboratory‐based dual stable isotope tracer approach (Na¹⁵NO₃ and NaH¹³CO₃), we evaluated the δ¹⁵N and δ¹³C isotope turnover rates in full‐grown adult invasive Limnomysis benedeni from Lake Constance. We provide δ¹⁵N and δ¹³C turnover rates based on nonlinear least‐squares regression and posterior linear regression models. Model precisions and fit were evaluated using Akaike's information criterion. Within a couple of days, the δ¹⁵N and δ¹³C of mysids began to change. Nevertheless, after about 14 days, L. benedeni did not reach equilibrium with their new isotope values. Since the experiment was conducted on adult subjects, it is evident that turnover was mainly influenced by metabolism (in contrast to growth). Unlike traditional dietary shifts, our laboratory‐based dual stable isotope tracer approach does not shift the experimental organisms into a new diet and avoids dietary effects on isotope values. Results confirm the application of isotopic tracers to label mysid subpopulations and could be used to reflect assimilation and turnover from the labeled dietary sources. Field‐based stable isotope studies often use isotopic mixing models commonly assuming diet‐tissue steady state. Unfortunately, in cases where the isotopic composition of the animal is not in equilibrium with its diet, this can lead to highly misleading conclusions. Thus, our laboratory‐based isotopic incorporation rates assist interpretation of the isotopic values from the field and provide a foundation for future research into using isotopic tracers to investigate invasion ecology.     

Water uptake by plants: perspectives from stable isotope composition

Stable isotope studies of hydrogen and oxygen stable isotope ratios of water within plants are providing new information on water sources, competitive interactions and water use patterns under natural conditions. Variation in the utilization of summer rain by aridland species and limited use of stream water by mature riparian trees are two examples of how stable isotope studies have modified our understanding of plant water relations. Analyses of xylem sap and tree rings have the potential of providing both short-term and long-term information on plant water use patterns.     

天山林区4种主要灌木夏季水分来源差异

天山林区群落结构相对简单、木本植物种类较少,但天山林区灌木群落中主要木本植物间的水分竞争模式尚不明确,水分利用动态缺乏定量分析。运用稳定同位素技术,对天山林区灌木群落4种主要灌木的茎杆水分及各潜在水源的氢氧稳定同位素组成进行测定,运用IsoSource模型定量分析4种灌木在夏季对各潜在水源的相对利用比例,探讨天山林区灌木群落主要灌木树种水分来源差异及动态变化。结果发现:7月,当浅层土壤含水量充足时,密刺蔷薇、黑果栒子和金丝桃叶绣线菊均大幅度吸收利用浅层土壤水,相对利用比例高于89.3%,异果小檗则相反,即吸收利用各潜在水源(浅层土壤水30.7%、中层土壤水29.4%、深层土壤水25.7%、溪水14.2%,下同);8月,当浅层土壤含水量降低时,密刺蔷薇转移至60-100 cm深层土壤水和溪水,相对利用比例分别为64.8%和27%,黑果栒子和金丝桃叶绣线菊以相似比例吸收利用各潜在水源(33.8%和36.8%、30.9%和29.7%、23.5%和22.3%、11.8%和11.2%),异果小檗则表现出可能吸收利用80-100 cm以下更稳定的深层土壤水;9月,当浅层土壤含水量升高时,4种灌木均大量吸收利用浅层土壤水,相对利用比例高于72.2%。这表明,天山林区灌木群落主要树种可通过可塑性转换水分来源来应对环境水分变异,在时间和空间上有效分割灌丛水源从而减缓对水分资源的竞争压力,从而通过在水分资源利用上的生态位分化促进物种间的共存。     

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(δS)＞枝条水氧同位素...     

Explaining geographical variation in the isotope composition of mouse lemurs (Microcebus)

Aim We sought to quantify geographical variation in the stable isotope values of mouse lemurs (Microcebus) and to determine whether this variation reflects trophic differences among populations or baseline isotopic differences among habitats. If the latter pattern is demonstrated, then Microcebus can become a proxy for tracking baseline habitat isotopic variability. Establishing such a baseline is crucial for identifying niche partitioning in modern and ancient communities. Location We studied five species of Microcebus from eight distinct habitats across Madagascar. Methods We compared isotopic variation in C₃ plants and Microcebus fur within and among localities. We predicted that carbon and nitrogen isotope values of Microcebus should: (1) vary as a function of abiotic variables such as rainfall and temperature, and (2) covary with isotopic values in plants. We checked for trophic differences among Microcebus populations by comparing the average difference between mouse lemur and plant isotope values for each locality. We then used multiple regression models to explain spatial isotope variation in mouse lemurs, testing a suite of explanatory abiotic variables. Results We found substantial isotopic variation geographically. Ranges for mean isotope values were similar for both Microcebus and plants across localities (carbon 3.5–4.0‰; nitrogen 10.5–11.0‰). Mean mouse lemur and plant isotope values were lowest in cool, moist localities and highest in hot, dry localities. Rainfall explained 58% of the variation in Microcebus carbon isotope values, and mean plant nitrogen isotope values explained 99.7% of the variation in Microcebus nitrogen isotope values. Average differences between mouse lemur and plant isotope values (carbon 5.0‰; nitrogen 5.9‰) were similar across localities. Main conclusions Isotopic data suggest that trophic differences among Microcebus populations were small. Carbon isotope values in mouse lemurs were negatively correlated with rainfall. Nitrogen isotope values in Microcebus and plants covaried. Such findings suggest that nitrogen isotope values for Microcebus are a particularly good proxy for tracking baseline isotopic differences among habitats. Our results will facilitate future comparative research on modern mouse lemur communities, and ecological interpretations of extinct Holocene communities.     

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。