O值和Keeling Plot方法区分麦田蒸散组分

利用稳定同位素技术和Keeling Plot方法可以有效分割地表蒸散量, 进而加深对陆地生态系统水循环的理解。该研究通过原位连续测定麦田的水汽同位素数据, 评价Keeling Plot方法在分割地表蒸散中的应用, 并揭示华北冬小麦(Triticum aestivum)蒸腾在总蒸散中的比例。实验于2008年3–5月在中国科学院栾城农业生态站进行, 利用国际上先进的H₂¹⁸O、HD¹⁶O激光痕量气体分析仪(TDLAS)为基础构建的大气水汽¹⁸O/¹⁶O和D/H同位素比原位连续观测系统, 同时利用涡度相关技术、真空抽提技术、同位素质谱仪技术, 获取了必要的数据。研究分析了一天中不同时间段的连续的大气水汽δ¹⁸O与水汽浓度倒数拟合Keeling Plot曲线的差异和可能的原因。结果显示, 中午时段的拟合结果较好, 这也暗示中午时段蒸腾速率高时最可能满足植物蒸腾的同位素稳定态假设。进一步的分析发现植物蒸腾的同位素稳定态并不总是成立, 尤其是水分胁迫下进入成熟期的小麦, 其蒸腾水汽同位素一般处于非稳定态。利用同位素分割结果显示, 生长盛期麦田94%–99%的蒸散来源于植物蒸腾。     

利用原位连续测定水汽δ~(18)O值和Keeling Plot方法 区分麦田蒸散组分

利用稳定同位素技术和Keeling Plot方法可以有效分割地表蒸散量,进而加深对陆地生态系统水循环的理解.该研究通过原位连续测定麦田的水汽同位素数据,评价Keeling Plot方法在分割地表蒸散中的应用,并揭示华北冬小麦(Triticum aes-tivum)蒸腾在总蒸散中的比例.实验于2008年3-5月在中国科学院栾城农业生态站进行,利用国际上先进的H_2~(18)O、HD~(16)O激光痕量气体分析仪(TDLAS)为基础构建的大气水汽~(18)O/~(16)O和D/H同位素比原位连续观测系统,同时利用涡度相关技术、真空抽提技术、同位素质谱仪技术,获取了必要的数据.研究分析了一天中不同时间段的连续的大气水汽δ~(18)O与水汽浓度倒数拟合Keeling Plot曲线的差异和可能的原因.结果显示,中午时段的拟合结果较好,这也暗示中午时段蒸腾速率高时最可能满足植物蒸腾的同位素稳定态假设.进一步的分析发现植物蒸腾的同位素稳定态并不总是成立,尤其是水分胁迫下进入成熟期的小麦,其蒸腾水汽同位素一般处于非稳定态.利用同位素分割结果显示,生长盛期麦田94%-99%的蒸散来源于植物蒸腾.     

华北低丘山区栓皮栎生态系统氧同位素日变化及蒸散定量区分

利用稳定同位素技术对华北低丘山区栓皮栎生态系统氧同位素日变化及蒸散定量区分进行研究,为华北低丘山区森林生态系统水汽交换研究提供基础。试验采用离轴积分腔输出光谱技术(OA-ICOS)连续测定生态系统不同高度水汽浓度和δ18O值,同时采用真空提取和液态水同位素分析仪测定枝条和土壤的δ18O值。结果显示,4个晴天中大气水汽浓度日变化复杂,变化趋势差异大,而δ18O日变化均成高-低-高的"V"型变化,最小值出现在12:00—18:00。Keeling方程在10:00—12:00的相关系数R2均大于0.71,方程达到极显著水平,表明此时段蒸腾速率较高,满足植物蒸腾的同位素稳定态假设。利用Keeling方程估算的栓皮栎生态系统δET值有相似的低-高-低日变化,与大气的δv值变化趋势相反。同位素分割结果显示栓皮栎生态系统中蒸腾占蒸散比例日变化呈现低-高-低的趋势,10:00—14:00蒸腾占蒸散比例达到90%以上,尽管6:00—10:00和14:00—18:00的蒸腾占蒸散比例下降,但平均值仍高达69.38%,表明华北低丘山区栓皮栎生态系统的蒸散主要来源于植物蒸腾。     

华北平原农田土壤蒸发δ18O的日变化特征及其影响因素

 土壤蒸发δ¹⁸O (δ_E)是影响大气水汽δ¹⁸O (δ_v)变异的重要因素, 也是农田生态系统蒸散组分土壤蒸发和植物蒸腾拆分的核心科学问题之一。δ_E主要基于Craig-Gordon模型计算, 主要受地表大气水汽δ_v、相对湿度(h)、平衡和动力学分馏系数以及土壤蒸发前缘液态水δ¹⁸O (δ_s)的影响。该研究以华北平原冬小麦(Triticum aestivum)-夏玉米(Zea mays)生态系统大气水汽δ_v的原位连续观测数据为基础, 同时结合不同深度的土壤日变化采样, 综合探讨了δ_E的日变化特征及其影响因素。结果表明: 冬小麦和夏玉米生长季δ_E的日变化表现为双峰曲线, 分别在6:00和15:00左右达到峰值。h强烈影响农田生态系统δ_E, 特别是在h > 95%的高相对湿度环境条件下Craig-Gordon模型并不适用。大气水汽δ_v的原位连续观测技术克服了传统的降水平衡预测大气水汽δ_v方法的不确定性, 可以显著提高δ_E的准确性。不同的平衡分馏系数对δ_E的结果无显著影响。不同的动力分馏系数尤其是考虑湍流扩散对动力分馏系数的影响会显著影响δ_E的模拟结果。土壤蒸发前缘的确定直接影响δ_s和标准化到土壤蒸发前缘温度下的h, 显著影响δ_E的准确性。结合动态箱或静态箱与稳定同位素红外光谱连续观测技术直接测定δ_E, 从而避免模型参数化过程引入的不确定性是未来研究的重要方向。     

叶片水H218O富集的研究进展

 植物叶片水H₂¹⁸O富集对大气中O₂和CO₂的¹⁸O收支有着重要影响。蒸腾作用使植物叶片水H₂¹⁸O富集, 而植物叶片水H₂¹⁸O富集的程度主 要受大气水汽δ¹⁸O和植物蒸腾水汽δ¹⁸O的影响。过去, 通过引入稳态假设(蒸腾δ¹⁸O等于茎水δ¹⁸O)得到Craig-Gordon模型的闭合形式, 或 将植物整个叶片水δ¹⁸O经过Péclet效应校正后得到植物叶片水δ¹⁸O的富集程度。然而, 在几分钟到几小时的短时间尺度上, 植物叶片蒸腾 δ¹⁸O是变化的, 稳态假设是无法满足的。最近成功地实现了对大气水汽δ¹⁸O和δD的原位连续观测, 观测精度(小时尺度)可达到甚至优于稳定 同位素质谱仪的观测精度。在非破坏性条件下, 高时间分辨率和连续的大气水汽δ¹⁸O和蒸腾δ¹⁸O的动态观测, 将提高植物叶片水H₂¹⁸O富集的 预测能力。该文综述了植物叶片水H₂¹⁸O富集的理论研究的新进展、研究焦点和观测方法所存在的问题, 旨在进一步加深理解植物叶片水H₂¹⁸O 富集的过程及其机制。     

叶片水H2^18O富集的研究进展

植物叶片水H2^18O富集对大气中O2和CO2的^18O收支有着重要影响。蒸腾作用使植物叶片水H2^18O富集,而植物叶片水H2^18O富集的程度主要受大气水汽δ^18O和植物蒸腾水汽δ^18O的影响。过去,通过引入稳态假设（蒸腾δ^18O等于茎水δ^18O）得到Craig-Gordon模型的闭合形式,或将植物整个叶片水δ^18O经过Peclet效应校正后得到植物叶片水δ^18O的富集程度。然而,在几分钟到几小时的短时间尺度上,植物叶片蒸腾δ^18O是变化的,稳态假设是无法满足的。最近成功地实现了对大气水汽δ^18O和δD的原位连续观测,观测精度（小时尺度）可达到甚至优于稳定同位素质谱仪的观测精度。在非破坏性条件下,高时间分辨率和连续的大气水汽δ^18O和蒸腾δ^18O的动态观测,将提高植物叶片水H2^18O富集的预测能力。该文综述了植物叶片水H2^18O富集的理论研究的新进展、研究焦点和观测方法所存在的问题,旨在进一步加深理解植物叶片水H2^18O富集的过程及其机制。     

黄土高原苹果园蒸腾导度大气驱动规律比较

植物蒸腾导度是表征土壤-植物-大气连续体（SPAC）中植物-大气间水汽传导过程、反映植物水分调控能力的一类重要变量,常见有冠层导度（G_c）、冠层气孔导度（G_s）与叶片气孔导度（g_s）,明确三者在反映冠层蒸腾过程时的异同或关联性对于理解植物水分利用机制具有重要意义。本研究基于对黄土高原果园苹果树生长季内树干液流（J_s）及环境因子的连续观测,计算了G_c、G_s及脱耦联系数（Ω）等变量,并与短期连续观测的叶片气孔导度（g_s）比较,分析了G_c、G_s和g_s在反映冠层蒸腾特征方面的异同及其关系。结果表明,日变化过程中G_s、g_s呈"单峰"型曲线,而G_c则呈"先增后减,午后抬升"的"双峰"型曲线。g_s与G_s存在较紧密的线性关系（R²=0.80）,但与G_c的线性关系较弱（R²=0.02）。G_c、G_s均随大气水汽压亏缺（VPD）的变化呈现确定的规律,其中,上边界函数呈递减的对数函数关系,平均值则符合先增后减的Log-Normal函数关系（R²>0.95）,拐点对应的VPD值分别为1.33和1.16 kPa。在一日内,G_s对VPD变化的响应过程与g_s对VPDL （基于叶片温度计算的水汽压亏缺）变化的响应过程总体一致,其一致性高于G_c对VPD变化的响应。整个生长季（4-10月）中果树的Ω平均值为0.12,随着Ω递减,G_c与G_s的线性相关性愈趋紧密,其斜率呈递增趋势,G_c越来越趋近于G_s。研究结果表明,在北方地区,基于树干液流的监测能较准确的推导整株并估算林分的冠层蒸腾导度。与实测g_s的变化过程比较,G_s比G_c具有更高的一致性,G_s可以作为描述苹果树水分利用过程响应大气驱动的更为恰当的变量。     

中国北方农牧交错带C3草本植物δ13C与温度的关系及其对水分利用效率的指示

由于植物稳定碳同位素组成(δ¹³C)综合反映了植物光合过程中C、H₂O交换的信息,因而从理论上讲它可以作为植物长期水分利用效率的潜在指标,并揭示与植物生理生态过程相联系的一系列气候环境信息。通过对中国北方农牧交错带400 mm等降水样带上28个科的118种C3草本植物叶片δ¹³C值的测定,探讨了C3草本植物δ¹³C和水分利用效率(WUE)对环境温度梯度变化的响应以及气候环境因素对其产生的影响,揭示了样带控制植物δ¹³C变化的主要环境因子。结果显示: 在400 mm降水带上C3植物δ¹³C分布区间为-31.5‰ -23.0‰,平均值为-27.6‰,分布范围与黄土区干旱-半干旱区C3草本植物一致。整体C3植物δ¹³C随年均温度和夏季均温升高分别变重0.14‰/℃和0.27‰/℃,指示植物WUE随大气温度升高而增加。但这仅是一种表象,温度与植物碳同位素的这种关系实质上是温度升高导致的土壤相对湿度(或湿润指数)降低造成水分胁迫进而影响植物碳同位素分馏的结果,植物可利用的有效水分是本样带植物碳同位素分馏的控制因子。5种C3广适性植物δ¹³C均随温度升高而变重,但变化幅度不同,而且它们之间平均δ¹³C值有显著差异,表明不同物种的水分利用状况对温度的响应不同,说明不同物种有不同适应环境变化的策略。此外,本结果还显示不同寿命的草本植物δ¹³C值以及用碳同位素表征的WUE表现出多年生草本>2年生草本>1年生草本(可能与不同寿命草本植物的根系分布和吸水能力有关),这与Ehleringer等在沙漠地区研究的结果一致,而与湿润气候区的结果相反,表明不同寿命的草本植物δ¹³C值和WUE的变化可能与当地水分条件有关。     

早春短命植物鸢尾蒜和准噶尔鸢尾蒜的光合途径

该研究以鸢尾蒜属两种早春短命植物准噶尔鸢尾蒜(Ixiolirion songaricum)和鸢尾蒜(Ixiolirion tataricum)为对象,通过解剖结构、光合参数、稳定碳同位素比值(δ¹³C)和光合关键酶活性分析二者的光合途径。结果发现：(1)显微结构和超微结构显示,两种短命植物的叶脉维管束鞘细胞1层、排列较紧密,鞘细胞内含有较多叶绿体且多离心分布,类似C₄花环结构。(2)准噶尔鸢尾蒜和鸢尾蒜最大净光合速率分别为14.81和15.04 μmol·m^-2·s^-1;两者的CO₂补偿点较低,分别为3.57和2.54 μmol·mol^-1,δ¹³C分别为-25.36±0.55‰和-25.76±1.38‰,光合酶PEPC/Rubisco比值分为别0.244和0.322。(3)最大净光合速率、δ¹³C和PEPC/Rubisco比值均说明两种植物为C₃光合途径,但二者均具有类似C₄的维管束鞘结构、较低的CO₂补偿点和暗呼吸速率,并具有高于部分C₃和C₃ C₄中间型植物的PEPC/Rubisco比值,表明两种短命植物的光合途径并非典型的C₃途径,而是C₃ C₄中间型。     

隧道工程对喀斯特槽谷区坡面产流及土壤侵蚀的影响

隧道工程导致地下水系统被破坏,但由此可能带来的土壤侵蚀却很少被涉猎。在重庆观音峡背斜隧道密集影响区和非隧道影响区的两个相邻小流域建立径流小区,基于高分辨率水文数据结合δD-H₂O、δ¹⁸O-H₂O同位素,对比两径流小区坡面、壤中产流规律和地表侵蚀产沙特征。结果表明,观测年内隧道影响区坡面产流对降雨响应更快,地表径流系数0.027,侵蚀模数16.68 t km^-2 a^-1;非隧道影响区地表径流系数0.013,侵蚀模数7.73 t km^-2 a^-1。相反,隧道影响区产生的壤中流产流系数仅为非隧道影响区的31%。对一场强降雨监测发现,两径流小区坡面流中δD-H₂O、δ¹⁸O-H₂O相似,但壤中流中却差异较大。用氢氧同位素混合模型分析得出隧道影响区坡面流、壤中流中降雨贡献率均大于非隧道影响区,侵蚀能力更强。这与土壤含水率减小和土壤结构的差异有关：隧道影响区土壤中粘粒的含量高于非隧道影响区,且出现上粘下松的异常土壤结构,使土壤下渗能力降低,地表径流增加。较小的土壤含水率与土壤粒径也有利于土壤搬运。研究为隧道工程导致的喀斯特区水土流失研究提供了基础数据,为喀斯特区水土流失防治和石漠化治理研究提供了新视角。     

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。     

Partitioning evapotranspiration fluxes from a Colorado grassland using stable isotopes: Seasonal variations and ecosystem implications of elevated atmospheric CO2

The stable isotopic composition of soil water is controlled by precipitation inputs, antecedent conditions, and evaporative losses. Because transpiration does not fractionate soil water isotopes, the relative proportions of evaporation and transpiration can be estimated using a simple isotopic mass balance approach. At our site in the shortgrass steppe in semi-arid northeastern Colorado, ¹⁸O values of soil water were almost always more enriched than those of precipitation inputs, owing to evaporative losses. The proportion of water lost by evaporation (E/ET) during the growing season ranged from nil to about 40% (to >90% in the dormant season), and was related to the timing of precipitation inputs. The sum of transpiration plus evaporation losses estimated by isotopic mass balance were similar to actual evapotranspiration measured from a nearby Bowen ratio system. We also investigated the evapotranspiration response of this mixed C₃/C₄ grassland to doubled atmospheric [CO₂] using Open-Top Chambers (OTC). Elevated atmospheric [CO₂] led to increased soil-water conservation via reduced stomatal conductance, despite greater biomass growth. We used a non-invasive method to measure the ¹⁸O of soil CO₂ as a proxy for soil water, after establishing a strong relationship between ¹⁸O of soil CO₂ from non-chambered control (NC) plots and ¹⁸O of soil–water from an adjacent area of native grassland. Soil–CO₂ ¹⁸O values showed significant treatment effects, particularly during a dry summer: values in ambient chambers (AC) were more enriched than in NC and elevated chamber (EC) plots. During the dry growing season of 2000, transpiration from the EC treatment was higher than from AC and lower than from NC treatments, but during 2001, transpiration was similar on all three treatments. Slightly higher evaporation rates from AC than either EC or NC treatments in 2000 may have resulted from increased convection across the soil surface from the OTC blowers, combined with lower biomass and litter cover on the AC treatment. Transpiration-use efficiency, or the amount of above-ground biomass produced per mm water transpired, was always greatest on EC and lowest on NC treatments.     

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     

Isotopic composition of transpiration and rates of change in leaf water isotopologue storage in response to environmental variables

During daylight hours, the isotope composition of leaf water generally approximates steady‐state leaf water isotope enrichment model predictions. However, until very recently there was little direct confirmation that isotopic steady‐state (ISS) transpiration in fact exists. Using isotope ratio infrared spectroscopy (IRIS) and leaf gas exchange systems we evaluated the isotope composition of transpiration and the rate of change in leaf water isotopologue storage (isostorage) when leaves were exposed to variable environments. In doing so, we developed a method for controlling the absolute humidity entering the gas exchange cuvette for a wide range of concentrations without changing the isotope composition of water vapour. The measurement system allowed estimation of ¹⁸O enrichment both at the evaporation site and for bulk leaf water, in the steady state and the non‐steady state. We show that non–steady‐state effects dominate the transpiration isoflux even when leaves are at physiological steady state. Our results suggest that a variable environment likely prevents ISS transpiration from being achieved and that this effect may be exacerbated by lengthy leaf water turnover times due to high leaf water contents.     

The use of stable isotopes to partition evapotranspiration fluxes into evaporation and transpiration

It is crucial to partition evapotranspiration (ET) into evaporation (E) and transpiration (T) components for better understanding eco-hydrological processes and their underlying mechanisms, and improving the establishment and validation of hydrological models at the ecosystem scale. Traditional eddy covariance technique serves as a useful tool to estimate ET, but it encounters difficulties in quantifying the relative contribution of E and T to ET. Combining with eddy covariance technique, it is possible to partition ET based on the measurements of stable oxygen and hydrogen isotopes in liquid and vapor phases of water in the Soil–Plant–Atmosphere Continuum (SPAC) system. The key challenge is to precisely determine the oxygen-18 and deuterium isotopic compositions of ET (δ_ET), E (δ_E) and T (δ_T). δ_E can be estimated based on the Craig–Gordon model. δ_T is usually approximated by the δ¹⁸O and δD of water in xylem or twig (δx), assuming δ_T equals δx under isotopic steady state (SSA). However, the SSA is only likely satisfied during midday in field conditions. The diurnal variations of δ_T is affected by isotopic composition of atmospheric water vapor (δ_v) and leaf water at the evaporating sites (δ_L,e), and relative humidity, resulting in the non-steady-state behavior of δ_T at the sub-daily cycles. δ_ET can be estimated using the flux-gradient approach or the Keeling plot by measuring the vapor mixing ratio and δ_v at different heights in the surface layer. However, δ_v observations by the traditional cold trap/mass spectrometer method are limited to a coarse time resolution, leading to discrete time series of δ_ET. It is now possible to make in situ and high time resolution measurements of δ_v and to analyze a large number of plant and soil samples due to technical and instrumental advances in recent years. It provides an opportunity to improve the model prediction of δ_L,e, and more importantly, to calculate δ_T from δ_L,e without invoking the SSA. Combining with the flux-gradient approach or the Keeling plot technique, continuous δ_ET measurements can be made. It offers us a premise for accurate ET partitioning on diurnal time scale. In this review we introduced the recent advances, foci and challenges for studies on ET partitioning using the stable isotopes technique.     

Contributions of evaporation, isotopic non-steady state transpiration and atmospheric mixing on the delta18O of water vapour in Pacific Northwest coniferous forests

Changes in the ²H and ¹⁸O of atmospheric water vapour provide information for integrating aspects of gas exchange within forest canopies. In this study, we show that diurnal fluctuations in the oxygen isotope ratio (δ¹⁸O) as high as 4‰ were observed for water vapour (δ¹⁸O_vp) above and within an old‐growth coniferous forest in the Pacific Northwest region of the United States. Values of δ¹⁸O_vp decreased in the morning, reached a minimum at midday, and recovered to early‐morning values in the late afternoon, creating a nearly symmetrical diurnal pattern for two consecutive summer days. A mass balance budget was derived and assessed for the ¹⁸O of canopy water vapour over a 2‐d period by considering the ¹⁸O‐isoflux of canopy transpiration, soil evaporation and the air entering the canopy column. The budget was used to address two questions: (1) do δ¹⁸O values of canopy water vapour reflect the biospheric influence, or are such signals swamped by atmospheric mixing? and (2) what mechanisms drive temporal variations of δ¹⁸O_vp? Model calculations show that the entry of air into the canopy column resulted in an isotopically depleted ¹⁸O‐isoflux in the morning of day 1, causing values of δ¹⁸O_vp to decrease. An isotopically enriched ¹⁸O‐isoflux resulting from transpiration then offset this decreased δ¹⁸O_vp later during the day. Contributions of ¹⁸O‐isoflux from soil evaporation were relatively small on day 1 but were more significant on day 2, despite the small H₂¹⁶O fluxes. From measurements of leaf water volume and sapflux, we determined the turnover time of leaf water in the needles of Douglas‐fir trees as ≈ 11 h at midday. Such an extended turnover time suggests that transpiration may not have occurred at the commonly assumed isotopic steady state. We tested a non‐steady state model for predicting δ¹⁸O of leaf water. Our model calculations show that assuming isotopic steady state increased isoflux of transpiration. The impact of this increase on the modelled δ ¹⁸O_vp was clearly detectable, suggesting the importance of considering isotopic non‐steady state of transpiration in studies of forest ¹⁸O water balance.     

Strong seasonal disequilibrium measured between the oxygen isotope signals of leaf and soil CO2 exchange

The oxygen isotope composition (δ¹⁸O) of atmospheric CO₂ is among a very limited number of tools available to constrain estimates of the biospheric gross CO₂ fluxes, photosynthesis and respiration at large scales. However, the accuracy of the partitioning strongly depends on the extent of isotopic disequilibrium between the signals carried by these two gross fluxes. Chamber‐based field measurements of total CO₂ and CO¹⁸O fluxes from foliage and soil can help evaluate and refine our models of isotopic fractionation by plants and soils and validate the extent and pattern of isotopic disequilibrium within terrestrial ecosystems. Owing to sampling limitations in the past, such measurements have been very rare and covered only a few days. In this study, we coupled automated branch and soil chambers with tuneable diode laser absorption spectroscopy techniques to continuously capture the δ¹⁸O signals of foliage and soil CO₂ exchange in a Pinus pinaster Aït forest in France. Over the growing season, we observed a seasonally persistent isotopic disequilibrium between the δ¹⁸O signatures of net CO₂ fluxes from leaves and soils, except during rain events when the isotopic imbalance became temporarily weaker. Variations in the δ¹⁸O of CO₂ exchanged between leaves, soil and the atmosphere were well explained by theory describing changes in the oxygen isotope composition of ecosystem water pools in response to changes in leaf transpiration and soil evaporation.     

The “isohydric trap”: A proposed feedback between water shortage,stomatal regulation,and nutrient acquisition drives differential growth and survival of European pines under climatic dryness

Climatic dryness imposes limitations on vascular plant growth by reducing stomatal conductance, thereby decreasing CO₂ uptake and transpiration. Given that transpiration‐driven water flow is required for nutrient uptake, climatic stress‐induced nutrient deficit could be a key mechanism for decreased plant performance under prolonged drought. We propose the existence of an “isohydric trap,” a dryness‐induced detrimental feedback leading to nutrient deficit and stoichiometry imbalance in strict isohydric species. We tested this framework in a common garden experiment with 840 individuals of four ecologically contrasting European pines (Pinus halepensis, P. nigra, P. sylvestris, and P. uncinata) at a site with high temperature and low soil water availability. We measured growth, survival, photochemical efficiency, stem water potentials, leaf isotopic composition (δ¹³C, δ¹⁸O), and nutrient concentrations (C, N, P, K, Zn, Cu). After 2 years, the Mediterranean species Pinus halepensis showed lower δ¹⁸O and higher δ¹³C values than the other species, indicating higher time‐integrated transpiration and water‐use efficiency (WUE), along with lower predawn and midday water potentials, higher photochemical efficiency, higher leaf P, and K concentrations, more balanced N:P and N:K ratios, and much greater dry‐biomass (up to 63‐fold) and survival (100%). Conversely, the more mesic mountain pine species showed higher leaf δ¹⁸O and lower δ¹³C, indicating lower transpiration and WUE, higher water potentials, severe P and K deficiencies and N:P and N:K imbalances, and poorer photochemical efficiency, growth, and survival. These results support our hypothesis that vascular plant species with tight stomatal regulation of transpiration can become trapped in a feedback cycle of nutrient deficit and imbalance that exacerbates the detrimental impacts of climatic dryness on performance. This overlooked feedback mechanism may hamper the ability of isohydric species to respond to ongoing global change, by aggravating the interactive impacts of stoichiometric imbalance and water stress caused by anthropogenic N deposition and hotter droughts, respectively.     

Non-steady state effects in diurnal 180 discrimination by Picea sitchensis branches in the field

We report diurnal variations in 18O discrimination (18 delta) during photosynthesis (18 delta A) and respiration (18 delta R) of Picea sitchensis branches measured in branch chambers in the field. These observations were compared with predicted 18 delta (18 delta pred) based on concurrent measurements of branch gas exchange to evaluate steady state and non-steady state (NSS) models of foliage water 18O enrichment for predicting the impact of this ecosystem on the Delta 18O of atmospheric CO2. The non-steady state approach substantially improved the agreement between 18 delta pred and observed 18 delta (18 delta obs) compared with the assumption of isotopic steady state (ISS) for the Delta 18O signature of foliage water. In addition, we found direct observational evidence for NSS effects: extremely high apparent 18 delta values at dusk, dawn and during nocturnal respiration. Our experiments also show the importance of bidirectional foliage gas exchange at night (isotopic equilibration in addition to the net flux). Taken together, neglecting these effects leads to an underestimation of daily net canopy isofluxes from this forest by up to 30%. We expect NSS effects to be most pronounced in species with high specific leaf water content such as conifers and when stomata are open at night or when there is high relative humidity, and we suggest modifications to ecosystem and global models of delta 18O of CO2.