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。     

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δ18O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。     

长江三角洲地区雨水中NH4+-N/NO3--N和δ15NH4+值的变化

2003年6月至2005年7月,利用自行设计的雨水收集器对位于长江三角洲地区的常熟、南京和杭州3个观测点进行了全年性雨水观测,分析了雨水中NH₄⁺-N/NO₃^--N和铵态氮自然丰度(δ¹⁵NH₄⁺)值的变化.结果表明：研究区3个观测点雨水中NH₄⁺-N/NO₃^--N和δ¹⁵NH₄⁺值均呈相似的季节性变化规律,两者的规律性变化在以田间农事耕作为主的常熟观测点尤其明显,而位于市区的南京观测点和位于城乡结合部的杭州观测点的规律性次之;雨水中NH₄⁺-N/NO₃^--N的峰值出现在6月下旬到8月上旬,然后逐渐下降,冬季降到最低;雨水中δ¹⁵NH₄⁺值在6月下旬到8月中旬为负值,在8月下旬到11月中下旬为正值,12月至翌年3月又变为负值,5至6月中旬又转变为正值.雨水中NH₄⁺-N/NO₃^--N和δ¹⁵NH₄⁺值的季节变化与不同作物生育期间氮肥的施用、当地气候的季节性变化以及其他NH3释放源的NH3挥发有关（人和动物排泄物、氮污染水体及有机氮源中的氨挥发）,其对大气湿沉降中NH₄⁺的来源、形态组成及陆地不同NH₃排放源的强度具有明显的指示意义.     

C4荒漠植物猪毛菜与木本猪毛菜的叶片解剖结构及光合生理特征

 为了比较C₄荒漠植物猪毛菜(Salsola collina)和木本猪毛菜(S. arbuscula)的抗旱结构和适应环境的光合作用特征, 在二者混生的群落中, 选择代表性植株, 采集叶片进行叶片解剖结构分析, 在自然条件下测定了二者叶片的气体交换参数。研究结果表明：猪毛菜叶片具表皮毛, 具有更发达的薄壁贮水组织;木本猪毛菜叶片具有更厚的角质层, 表皮下有1层下皮细胞, 其栅栏组织细胞较长, 排列更紧密。猪毛菜的净光合速率明显高于木本猪毛菜, 日平均值分别为21.5和15.7 μmol CO₂·m^–2·s^–1。猪毛菜的蒸腾速率也明显高于木本猪毛菜, 日平均值分别为14.9和10.2 mmol·m^–2·s^–1。猪毛菜和木本猪毛菜的水分利用效率的日平均值分别为1.39和1.53 μmol CO₂·mmol^–1 H₂O, 特别是在14:00时分别为1.61和2.30 μmol CO₂·mmol^–1 H₂O, 木本猪毛菜高出猪毛菜约42%。猪毛菜的光补偿点低于木本猪毛菜, 而光饱和点和光量子效率较高, 具有更低的CO₂补偿点。这表明：二者的旱生结构不同, 木本猪毛菜具有更显著的荒漠植物特征;在适于二者混生的环境下, 猪毛菜比木本猪毛菜的光合能力更强, 而木本猪毛菜的水分利用效率更高。     

外源H2O2处理对唐古特白刺愈伤组织脯氨酸代谢的影响

以唐古特白刺（Nitraria tangutorum Bobr.）愈伤组织为材料,研究外源H₂O₂（2和10 μmol·L^-1）处理下其脯氨酸含量及相关代谢酶活性的变化,试图从细胞水平揭示H₂O₂影响脯氨酸代谢的生理机制。结果显示,2和10 μmol·L^-1 H₂O₂处理24 h使唐古特白刺愈伤组织脯氨酸含量分别变为对照的112%和92%,而处理72 h后,脯氨酸含量增加为对照的141%和119%;与对照相比,外源H₂O₂处理诱导愈伤组织脯氨酸脱氢酶活性降低,而谷氨酸激酶活性升高,但鸟氨酸转氨酶活性无显著变化;此外,H₂O₂处理使唐古特白刺愈伤组织内源性H₂O₂含量升高。结果表明,外源H₂O₂诱导了唐古特白刺愈伤组织H₂O₂含量的增高和脯氨酸的积累,且H₂O₂处理下脯氨酸脱氢酶活性的降低及谷氨酸激酶的升高与愈伤组织脯氨酸的积累有关。     

外源氮对沼泽湿地CH4和N2O通量的影响

三江平原沼泽湿地受到大气沉降、地表径流、农业排水等外源氮素的输入,对湿地生态系统CH₄和N₂O通量有重要影响。采用野外原位施肥试验模拟外源氮输入,设0,60,120,240kgN•hm^-2 4种试验处理,探讨外源氮对沼泽湿地CH₄和N₂O通量的影响。结果表明,外源氮促进了CH₄和N₂O排放。与对照处理比较,各施氮水平CH₄平均排放通量分别增加了181％,254％和155％,N2O排放通量分别增加了21％,100％和533％。外源氮输入对CH₄排放的季节变化形式影响不大,而N²O的季节变化形式随着氮输入表现出波动变化的趋势。不同施氮水平对CH₄排放的促进作用与植物生长阶段和产CH₄的微生物过程密切相关,N₂O排放通量随氮输入量呈指数增加（R²＝0.97,p<0.01）。外源氮通过影响湿地微生物过程来进一步影响CH₄和N₂O的排放。     

植物细胞H2O2的信号转导途径

光、环境胁迫和植物激素ABA可以引起植物体内H₂O₂升高, 而H₂O₂作为一个较早进化出来的信号分子, 不仅在诱导氧化性光合作用中起了关键的作用, 并且可以调节诸如气孔运动、超敏反应、细胞凋亡和基因表达等许多过程. 细胞内H₂O₂浓度必须维持在一种精细平衡状态, 它一方面可以通过质膜氧化还原系统和光呼吸系统产生, 另一方面也存在完善的清除机制. H₂O₂从质外体或者产生源进入细胞, 然后进入亚细胞区域. H₂O₂可以调节信号转导蛋白, 如蛋白质磷酸化酶、转录因子、以及位于质膜或其它膜上的Ca²⁺通道. 其中, 蛋白质可逆磷酸化可启动细胞质和细胞核的下游信号转导, 通过影响转录因子而影响基因的表达; 转录因子通过氧化而激活自身或诱导其定向转运至细胞核内. 然而, H₂O₂作为信号分子的研究相对处于“年轻”阶段, 诸如细胞如何感受H₂O₂, 以及在细胞感受H₂O₂信号转导过程中哪种细胞过程是最主要的或是限速步骤, 何种基因对H₂O₂是特异和必需的等问题仍然所知甚少, 这些问题的破解依赖于功能基因组学和遗传学分析.     

黑河中游春玉米叶片水δD和δ~(18)O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(δ~(18)O l,b)和氘(D)同位素比值(δDl,b)(合称δl,b)实测数量少只能作为模型验证数据,导致δl,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δl,b富集的控制机制研究。叶片水δDl,b和δ~(18)O l,b的富集程度(ΔDl,b和Δ18O l,b,合称Δl,b)通常表示为δl,b与茎秆水D同位素比值(δDx)和18O同位素比值(δ~(18)Ox)(合称δx)之差,即Δl,b=δl,b–δx。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δl,b和δx数据,配套开展了大气水汽δ~(18)O和δD(合称δv)等辅助变量的原位连续观测,探讨了季节和日尺度上的δl,b富集特征及其影响因素。结果表明:叶片水δl,b和Δl,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δv和相对湿度是δDl,b和ΔDl,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δ~(18)O 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ΔDl,b和Δ18Ol,b与影响因素的差异性,有助于理解叶片水δD和δ~(18)O富集过程以及对模型发展有一定的指导意义。     

Na2CO3胁迫下星星草幼苗活性氧及保护酶活性的变化

对低浓度Na₂CO₃胁迫下星星草幼苗相对电导率、O^-₂产生速率、H₂O₂含量以及保护酶CAT、SOD和POD活性的研究结果表明,低盐胁迫1 d后,星星草幼苗细胞膜的通透性、O^-₂产生速率、H₂O₂含量及保护酶活性都随着盐胁迫的加剧而升高,其具体的变化规律与盐胁迫强度和幼苗细胞膜的受损伤程度密切相关,但相关关系的性质上具有差异。     

Photoreduction of 18O2 and H218O2 with Concomitant Evolution of 16O2 in Intact Spinach Chloroplasts: Evidence for Scavenging of Hydrogen Peroxide by Peroxidase

Illuminated intact spinach chloroplasts decomposed one moleculeof H₂¹⁸O₂ which resulted in the evolution of a half moleculeof ¹⁶O₂, but little ¹⁸O₂. The chloroplasts showed the same rateof photoreduction of ¹⁸C₂ as that of the evolution of ¹⁶O₂ withoutaccumulation of H₂¹⁸O₂. These reactions were suppressed by DCMU,and also by several inhibitors of ascorbate peroxidase and dehydroascorbateand monodehydroascorbate reductases in chloroplasts. These observationsindicate that the hydrogen peroxide produced in chloroplastsis reduced to water by a peroxidase using a photoreductant asthe electron donor. The hydrogen peroxide scavenging systemof chloroplasts was inactivated if hydrogen peroxide was addedin the dark, but not if added during the light. (Received May 4, 1984; Accepted July 10, 1984)     

Diurnal variation of Δ13CO2, ΔC18O16O and evaporative site enrichment of δH218O in Piper aduncum under field conditions in Trinidad

Concurrent measurements of gas exchange, instantaneous isotope discrimination (Δ) against ¹³CO₂ and C¹⁸O¹⁶O, and extent of ¹⁸O enrichment in H₂O at the evaporative sites, were followed in a tropical forest pioneer, Piper aduncum, on two different days in Trinidad during February 1995. Δ¹³CO₂ differed from that predicted from measurements of internal:external CO₂ concentration (C_i/C_a) and showed a wide range of values which decreased throughout the course of the day. Derivation of C_c (the CO₂ concentration at the carboxylation site) was not possible using carbon isotope discrimination under field conditions in situ and was derived assuming a constant value of internal transfer conductance (g_w). Under low rates of assimilation the derived C_c/C_a, like C_i/C_a, remained relatively stable over the course of both days and ΔC¹⁸O¹⁶O followed evaporative demand. Lower values of ΔC¹⁸O¹⁶O on day 2 occurred in response to the indirect effect of increased leaf-to-air vapour pressure deficits (VPD) and reduced stomatal conductance. For the first time, direct determination of the δH₂¹⁸O of transpired water vapour (δ_t) allowed derivation of evaporative site enrichment without the prerequisite of isotopic steady state (ISS) defined in the Craig and Gordon model. Generally, δ_t was less enriched than the source water (δ_s) in the morning and more enriched in the afternoon, which would be predicted from an increase and decrease in ambient VPD, respectively. On both days, leaves of P. aduncum approached ISS (indicated where δ_t≈δ_s) between 1300 and 1500 h. Evaporative site enrichment was maintained into the late afternoon, despite a decrease in ambient VPD. The data presented provide a greater insight into the natural variation in isotopic discrimination under field conditions, which may help to refine models of terrestrial biome discrimination.     

准噶尔荒漠早春短命植物的光合特性及生物量分配特点

准噶尔荒漠分布的早春短命植物不仅具有十分独特的生物学特点,而且在荒漠植物群落演替、物种多样性维持及土壤改良与防治水土流失等方面具有重要的生态学价值。该文运用Li-6400开放式气体交换光合作用测定系统,对分布于准噶尔荒漠的16种早春短命植物生长盛期的净光合速率(P_n)、蒸腾速率(T_r)、水分利用效率(WUE)等特征进行了测定,并对其中7种植物与生长相关的生物量分配特征进行了分析。结果表明:1)16种植物的最大P_n、 最大T_r及WUE分别为8.07~35.96 μmol CO₂·m^-2·s^-1、3.16~29.64 mmol H₂O·m^-2·s^-1、0.54~4.26 μmol CO₂·mmol^-1H₂O;种间最大P_n与最大气孔导度(Stomatal conductance, G_s)之间存在正相关关系,其相关系数为0.77(p<0.05),线性回归斜率为26.36 μmol·mmol^-1;从光合速率对胞间CO₂浓度及光量子通量密度的响应曲线来看,这类植物的表观CO₂补偿点均在4~5 Pa之间(28~30 ℃),表观羧化效率为0.64~1.86 μmol CO₂·m^-2·s^-1·Pa^-1,表观量子效率为0.05~0.06。2)从生物量分配来看,所测植物的个体生物量为0.05~0.39 g;单株总叶面积为 3.24~51.40 cm²;单位叶面积干重为0.40~0.77 g·m^-2,根在总生物量中所占比例为5.72%~19.43%,单株叶面积比在2.92~9.00 m²·kg^-1之间。种间根所占生物量的比与对应的WUE之间的比较分析结果表明,二者之间存在显著的正相关关系,其相关系数r为0.93(p<0.01)。这些结果表明,所观测的早春短命植物具有典型的C₃植物特征,相比其它类型的荒漠植物具有较高的单位叶面积P_n、高T_r及低WUE,并且在生长发育过程中表现出很低的根/地上生物量比、较高的叶面积比和单位叶面积干重,说明它们具有相对高的生长速率,这与其生长发育节律相一致,反映了它们与准噶尔荒漠环境相适应的特点。     

Isotopic heterogeneity of water in transpiring leaves: identification of the component that controls the δ18O of atmospheric O2 and CO2

Two direct but independent approaches were developed to identify the average δ¹⁸O value of the water fraction in the chloroplasts of transpiring leaves. In the first approach, we used the δ¹⁸O value of CO₂ in isotopic equilibrium with leaf water to reconstruct the δ¹⁸O value of water in the chloroplasts. This method was based on the idea that the enzyme carbonic anhydrase facilitates isotopic equilibrium between CO₂ and H₂O predominantly in the chloroplasts, at a rate that is several orders of magnitude faster than the non-catalysed exchange in other leaf water fractions. In the second approach, we measured the δ¹⁸O value of O₂ from photosynthetic water oxidation in the chloroplasts of intact leaves. Since O₂ is produced from chloroplast water irreversibly and without discrimination, the δ¹⁸O value of the O₂ should be identical to that of chloroplast water. In intact, transpiring leaves of sunflower (Helianthus annuus cv. giant mammoth) under the experimental conditions used, the average δ¹⁸O value of chloroplasts water was displaced by 3—10 % (depending on relative humidity and atmospheric composition) below the value predicted by the conventional Craig & Gordon model. Furthermore, this δ¹⁸O value was always lower than the δ¹⁸O value that was measured for bulk leaf water. Our results have implications for a variety of environmental studies since it is the δ¹⁸O value of water in the chloroplasts that is the relevant quantity in considering terrestrial plants influence on the δ¹⁸O values of atmospheric CO₂ and O₂, as well as in influencing the δ¹⁸O of plant organic matter.     

Temporal and spatial variations in the oxygen-18 content of leaf water in different plant species

Temporal variations in the δ¹⁸ oxygen (δ¹⁸O) content of water transpired by leaves during a simulated diurnal cycle fluctuated around the δ¹⁸O content of the source water. Reconstructed variations in the δ¹⁸O values of leaf water differed markedly from those predicted by conventional models. Even when transpiring leaves were maintained under constant conditions for at least 3 h, strict isotopic steady-state conditions of leaf water (equality of the ¹⁸O/¹⁶O ratios in the input and transpired water) were rarely attained in a variety of plant species (Citrus reticu-lata, Citrus paradisi, Gossypium hirsutum, Helianthus annuns, Musa musaceae and Nicotinia tabacum). Isotopic analysis of water transpired by leaves indicated that leaves approach the isotopic steady state in two stages. The first stage takes 10 to 35 min (with a rate of change of about 3–3%h^?1), while in the second stage further approach to the isotopic steady state is asymptotic (with a rate of change of about 0–4% h^?1), and under conditions of low transpiration leaves can last for many hours. Substantial spatial isotopic heterogeneity was maintained even when leaves were at or near isotopic steady state. An underlying pattern in this isotopic heterogeneity is often discerned with increasing ¹⁸O/¹⁶O ratios from base to tip, and from the centre to the edges of the leaves. It is also shown that tissue water along these spatial isotopic gradients, as well as the average leaf water, can have ¹⁸O/¹⁶O ratios both lower and higher than those predicted by the conventional Craig and Gordon model. We concluded, first, that at any given time during the diurnal cycle of relative humidity the attainment of an isotopic steady state in leaf water cannot be assumed a priori and, secondly, that the isotopic enrichment pattern of leaf water reflects gradual enrichment along the water-flow pathway (e.g. as in a string of pools), rather than a single-step enrichment from source water, as is normally assumed.     

Effects of Different CO2 Environments on the Photosynthesis-Yield Relationship and the Carbon and Water Balance of a White Clover (Trifolium repens L. cv. Blanca) Sward

Effects of atmospheric CO₂ enrichment to a level above 600 parts10^–6 on leaf and canopy gas exchange characteristics wereinvestigated in Trifolium repens, using an open system for gasexchange measurement. The cuvettes of the system served as growthchambers, allowing continuous measurement in a semi-controlledenvironment of ±350 and ±600 parts 10^–6CO₂, respectively. Carbon balance data were compared with cropyield and effects on the canopy level were compared with measuredleaf responses of photosynthesis and stomatal behaviour. Photosyntheticstimulation by high CO₂ was stronger at the canopy level (103%on average) than for leaves (90% in full light), as a consequenceof accelerated foliage area development. The latter increasedabsolute water consumption by 16%, despite strong stomatal closure.The overall result was a 63% improvement in canopy water useefficiency (WUE), while leaf WVE increased almost 3-fold insaturating light. The stomatal response was such that, whilethe internal CO₂ concentration in the leaf, ch increased withrising atmospherical CO₂ concentration, c_a, ci/c_a was somewhatdecreased. Total canopy resistance, R_c, was generally lowerat high CO₂ levels, despite higher leaf resistance. Higher canopyCO₂ loss at night and faster light extinction in a larger-sizedhigh CO₂ canopy were major drawbacks which prevented a furtherincrease in dry matter production (the harvest index was increasedby a factor 1.83). Key words: CO₂ enrichment, canopy CO₂ exchange, carbon balance, water use efficiency, leaf and canopy resistance     

The Responses of Net Photosysthesis to Light and Temperature in Spartina townsendii(sensu lato), a C4 Species from a Cool Temperate Climate

Carbon dioxide and water vapour exchanges for single attachedleaves of the temperate C₄ grass Spartina townsendii were measuredunder controlled environment conditions in an open gas-exchangesystem. The responses of net photosynthesis, stomatal resistance,and residual resistance to leaf temperature and photon fluxdensity are described. The light and temperature responses ofnet photosynthesis in S. townsendii are compared to informationon these responses in both temperate C₃ grasses and sub-tropicalC₄ grasses. Adaptation of photosynthesis in this C₄ speciesto a cool temperate climate is indicated both by the light andtemperature responses of net photo-synthesis. Unlike the C₄grasses examined previously, significant rates of net photosynthesiscan be detected at leaf temperatures below 10?C. Rates of netphotosynthesis equal or exceed those reported for temperateC₃ grasses at all of the temperature (5–40?C) and photonflax density (13–2500µmol m^–2 s^–1) conditionsexamined. Maximum rates of net photosynthesis in S. townsendiiare almost double those reported for C₃ herbage grasses. Unliketemperate C₃ grasses, the major limitation to net photosynthesisat low leaf temperatures (10?C and below) is the stomatal resistance,showing that the low residual resistance characteristic of C₄species is maintained in S. townsendii even at low leaf temperatures.     

Modeling biophysical controls on canopy foliage water 18O enrichment in wheat and corn

Leaf water ¹⁸O enrichment is an important factor controlling the H₂¹⁸O, C¹⁸OO, and O¹⁸O exchanges between the biosphere and the atmosphere. At present, there is limited capacity to explain the enrichment mechanisms in field conditions. In this study, three models of varying complexity were used to simulate the leaf water ¹⁸O enrichment at the canopy scale. Comparisons were made among the models and with high‐frequency isotopic measurements of ecosystem water pools in wheat and corn. The results show that the steady state assumption was a better approximation for ecosystems with lower canopy resistance, that it is important to consider the effect of leaf water turnover in modeling the enrichment and not necessary to deal with time changes in leaf water content, and that the leaf‐scale Péclet effect was incompatible with the big‐leaf modeling framework for canopy‐air interactions. After turbulent diffusion has been accounted for in an apparent kinetic factor parameterization, the mean ¹⁸O composition of the canopy foliage water was a well‐behaved property predictable according to the principles established by leaf‐scale studies, despite substantial variations in the leaf water enrichment with leaf and canopy positions. In the online supplement we provided a discussion on the observed variability of leaf water ¹⁸O composition with leaf and canopy positions and on the procedure for correcting isotopic measurements for organic contamination.     

Correlating δ13C and δ18O in cellulose of trees

We measured the carbon and oxygen isotopic composition of stem cellulose of Pinus sylvestris, Picea abies, Fagus sylvatica and Fraxinus excelsior. Several sites along a transect of a small valley in Switzerland were selected which differ in soil moisture conditions. At every site, six trees per species were sampled, and a sample representing a mean value for the period from 1940 to 1990 was analysed. For all species, the mean site δ¹³C and δ¹⁸O of stem cellulose are related to the soil moisture availability, whereby higher isotope ratios are found at drier sites. This result is consistent with isotope fractionation models when assuming enhanced stomatal resistance (thus higher δ¹³C of incorporated carbon) and increased oxygen isotope enrichment in the leaf water (thus higher δ¹⁸O) at the dry sites. δ¹⁸ O-δ¹³C plots reveal a linear relationship between the carbon and oxygen isotopes in cellulose. To interpret this relationship we developed an equation which combines the above-mentioned fractionation models. An important new parameter is the degree to which the leaf water enrichment is reflected in the stem cellulose. In the combined model the slope of the δ¹⁸O-δ¹³C plot is related to the sensitivity of the p_i/p_a of a plant to changing relative humidity.     

Photosynthesis, Respiration, and Carbon Assimilation in Water-Stressed Maize at Two Oxygen Concentrations

Photosynthesis decreased with decreasing leaf water potentialas a consequence of stomatal closure and possibly non-stimataleffects of severe stress. Assimilation ceased at c. 16x 10⁵Pa. Photo-respiration, in 21% O₂, was small in relation to assimilationin unstressed leaves and decreased as leaf water potential fellbut it was much larger in proportion to photosynthesis at severestress. Decreasing the O₂ content to 1.5% increased photosynthesisslightly and decreased photo-respiration but did not changethe stress at which assimilation stoped. Dark respiration wasinsensitive to both O₂ and stress. Less ¹⁴C accumulated in stressedleaves but in 21% O₂ a greater proportion of it was in aminoacids, particularly glycine and serine. 1.5% O₂ decreased the¹⁴C in glycine to 10% and in serine to 50% of their levels in21% O₂. In both O₂ concentrations the proportion of ¹⁴C in serineincreased only at the most severe stress. Gas exchange measurementsand changes in the ¹⁴C flux to glycine are interpreted as theresult of glycolate pathway metabolism increasing as a proportionof assimilation in stressed leaves in high O₂. The small absoluterate of photorespiration in high O₂ and at low leaf water potentialmay be due to slow rates of glycine decarbodylation as wellas efficient fixation of any CO₂ produced. Serine is synthesizedby an O₂-sensitive pathway and an O₂-insensitive pathway, whichis most active at severe stress. Synthesis of alanine competeswith that of glycine and serine for a common precursor suppliedby the photo-synthetic carbon reduction cycle. The relativespecific radioactivities of aspartate and alanine suggest thatthey are derived from a common precursor pool, probably pyruvatefrom 3-PGA. The amounts of 3-PGA, aspartate, malate, alanine,and sucrose decreased with increasing water stress as a consequenceof slower assimilation and pool filling. Other amino acids,glycine, serine, glutamate, and proline, accumulated at lowwater potential possibly due to increased synthesis and slowerrates of consumption. Changes in pool sizes, carbon fludes,and specific activities of metabolites are related to the mechanismof C₄ photosynthesis and current concepts of glycolate pathwaymetabolism.