中国区域478种C3植物叶片碳稳定性同位素组成与水分利用效率

目前, 在中国区域关于植物碳稳定性同位素组成(δ¹³C)已经有了很多的研究, 同时, δ ¹³C作为植物水分利用效率(WUE)的替代指标, 得到了越来越广泛的应用。而这些研究多集中在站点或小的区域尺度, 那么在整个中国区域尺度上, δ¹³C能否作为植物WUE的替代指标值得探讨。该文通过对文献资料的收集整理, 研究了中国区域187个采样点478种C₃植物叶片的δ¹³C, 统计分析结果表明δ¹³C的变化范围为-33.50‰- -22.00‰, 均值为-(27.10 ± 1.70)‰。在乔木、灌木和草本3种不同的生活型间, 叶片δ¹³C的差异达到极显著水平, 其中以草本的δ¹³C最高, 乔木最低, 这与在站点或小的区域尺度上的研究结果不同。对不同系统发育类型的植物而言, 种子植物的δ¹³C极显著地大于蕨类植物; 祼子植物与被子植物间的差异未达到显著水平; 单子叶植物极显著地大于双子叶植物。叶片δ¹³C值随经度的变化没有明显的规律, 但是随纬度的增加, δ ¹³C极显著地升高。随年均温度和年均降雨量的降低, 叶片δ¹³C值极显著升高。年均降雨量与δ¹³C间的这种极显著的负相关关系, 与WUE和降水量间的关系一致, 这表明在大的区域尺度上, δ¹³C可以作为植物WUE的指示指标。     

马占相思叶片液汁碳同位素甄别率和水分利用效率

利用PMS压力室压取叶片液汁并借助稳定碳同位素质谱仪测定碳同位素比率,分析了马占相思(Acacia mangium )叶片液汁光合产物的甄别率(△)和估测了水分利用效率.结果表明:阴天马占相思林冠层日平均空气CO2 的碳同位素比率(δa)为-7.57‰±1.41‰,晴天则为-8.54‰±0.67‰;阴天叶片液汁的光合产物碳同位素比率(δp)日变化呈鞍型,而晴天则从早上至黄昏逐步降低;晴天δp与叶片/空气水汽压亏缺(D)明显呈负相关,阴天的δp变化相对较小;δp随叶片水势(ψ)降低而降低,显示水分胁迫引起δp降低;叶片液汁的D和经气体交换法获得的细胞胞间(Pi)和外界(Pa)CO2分压之比呈正相关,测定结果与理论上碳同位素相关扩散和生化分馏相一致.分析结果显示,田间马占相思空气CO2经气孔扩散的稳定同位素效应口a=4.6‰,有关Pi的净固定的稳定碳同位素效应6=28.2‰.认为液汁碳同位素甄别率是外围空气CO2进入光合产物的净甄别率,由叶片液汁△估测的水分利用效率与气体交换法所得结果相一致(R2=0.86,P＜0.001).本文所采用的叶片液汁光合产物测定即时△以及计算水分利用效率的方法,可减少田间条件下环境因素明显变化对水分和碳同位素的影响,该方法有助于进一步开展由植株至冠层扩展的碳和水分平衡的生理生态研究.     

植物叶片水分利用效率研究综述

植物能否适应当地的极限环境条件,最主要的看它们能否很好地协调碳同化和水分耗散之间的关系,即植物水分利用效率(WUE)是其生存的关键因子.就近来研究最多的叶片水平上的WUE,从叶片WUE的定义,方法,进展等方面对其进行总结概括,并就今后植物叶片水分利用效率的研究提出了几点看法:方法上,叶片碳同位素方法是目前植物叶片长期水分利用效率研究的最佳方法,而δ13C的替代指标将继续是方法研究中的一个方向,前景乐观;研究内容上,要加强极端干旱区河岸林木的δ13C和WUE的研究;结合植物生理生态学,生物学和稳定同位素技术,探究植物叶片长期水分利用效率的机理,特别是要加强运用双重同位素模型加深和理解植物叶片长期水分利用效率变化规律和内在机制的研究;要结合多种方法,加强多时空尺度植物叶片WUE及其之间的转换研究.     

干旱胁迫下四倍体刺槐幼苗水分利用效率及稳定碳同位素组成的研究

在模拟自然干旱的条件下,测定2个四倍体刺槐品种(K2、K3)和普通二倍体刺槐(K1)一年生组培苗的长期和瞬时水分利用效率(WUE)及其稳定碳同位素组成(δ13C),以探讨四倍体刺槐的抗旱机理.结果表明,K2、K3的长期水分利用效率(WUEL)在不同水分胁迫条件下都显著高于K1,它们在同等供水条件下比K1具有更大的生物量产出;3个材料叶片的瞬时水分利用效率(WUEI)均随干旱胁迫的加剧表现先升后降的趋势,且胁迫处理均高于适宜水分处理.随水分胁迫的加剧,各品种刺槐苗木叶片的δ13C显著升高;K2、K3的δ13C在各水分条件下均高于K1;各材料叶片的δ13C与其WUEL有良好的正相关性,可以作为筛选高WUE刺槐品种的指标.     

不同生活型绿化植物叶片碳同位素组成的季节特征

通过测定北京地区不同生活型绿化植物叶片的碳同位素组成(δ13C值),从植物种和生活型两个方面研究植物水分利用效率的自然可变性。结果表明:所测定的75种植物(隶属于35科65属)的叶片的δ13C值变幅,春季为-30.7‰--23.4‰,夏季为-31.5‰--25.1‰,秋季为-31.4‰--23.9‰;落叶灌木种间差异不显著(p=0.114),而常绿乔木(p=0.005)、落叶乔木(p0.001)、常绿灌木(p=0.022)、草本植物(p0.001)和藤本植物(p=0.001)的种间差异显著或极显著;同一生活型植物叶片的δ13C季节差异显著,春季叶片的δ13C值显著大于夏秋两季(常绿乔木除外),不同生活型植物叶片的δ13C值在春、夏、秋3个季节差异都达到了极显著水平(春季p=0.001、夏季p0.001、秋季p0.001),且叶片的δ13C值表现出乔木树种灌木树种藤本植物草本植物、常绿植物落叶植物的规律。因此,植物种和生活型均会引起植物叶片δ13C值的变化,但δ13C受生活型变化的影响较大,表明不同生活型植物的水分利用效率具有明显差异。     

广西弄岗喀斯特季节性雨林藤本种子植物多样性及繁殖习性

藤本植物是森林生物多样性的重要组成部分,但在生物多样性丰富的喀斯特森林中,藤本植物的组成和繁殖物候等特征鲜为人知。为此,作者通过长期的调查监测,对弄岗喀斯特季节性雨林藤本种子植物的种类组成、物种分布、开花和结实时间以及种实类型进行了整理和分析。结果表明:该区藤本种子植物种类丰富,共有333种,隶属56科145属,其中藤状灌木119种,草质藤本88种,木质藤本126种。不同生活型的藤本在不同地貌部位的种数分布有所差异,草质藤本较多分布在洼地,藤状灌木和木质藤本较多分布在坡地。藤本种子植物的开花结实表现出一定的季节性,开花高峰期在4–9月,结实高峰期在7–12月,其中藤状灌木的季节性相对较弱。藤本种子植物的开花比率与降水量和气温呈极显著的正相关关系,每年雨季即为藤本种子植物的开花高峰期。结实高峰期比开花高峰期滞后约3个月,出现在雨季末期或雨季结束后。在木质藤本中,翅果比率与风速极显著正相关,与降水量和气温极显著负相关,说明了翅果型木质藤本趋于在高风速的干旱季节结实。总之,弄岗喀斯特季节性雨林藤本种子植物的多样性和繁殖特征与生境资源的时空变异相关联。     

桂西北喀斯特坡地典型生境不同植物叶片的碳同位素差异

通过测定7种典型植物的叶片δ13C值,比较了桂西北喀斯特坡地连片裸露石丛及其临近浅薄土层处植物叶片δ13C值及其季节性差异。结果表明:旱季(2008年11月、2009年3月)植物叶片δ13C值显著高于雨季(2009年7月)(P0.05);旱季和雨季菜豆树、铁线莲、黄荆的δ13C值无显著差异(P0.05),而盐肤木、红背山麻杆、蕨、肾蕨旱季显著高于雨季(P0.05);季节性降雨差异对乔木类菜豆树、藤本类的铁线莲以及根系较深灌木叶片δ13C值的影响较小,而对浅根系草本植物、浅根灌木影响较大;旱季和雨季2种生境下菜豆树叶片δ13C值差异均不大,旱季和雨季连片裸岩生境的红背山麻杆和黄荆叶片δ13C值与临近浅薄土层生境值的差异虽不显著(P0.05),但差异大于菜豆树,说明生境条件的差异对这2种灌木叶片δ13C值的影响较大。     

滨海沙地不同树种人工林叶片和土壤表层稳定碳氮同位素及水分利用效率研究

以福州市滨海后沿沙地人工营造的湿地松、木麻黄、尾巨桉、肯氏相思和纹荚相思防护林为研究对象,测定不同年龄(新叶、老叶)叶片、表层土壤(0~10cm)天然稳定碳、氮同位素丰度值(δ~(13) C、δ~(15)N),研究稳定碳、氮同位素丰度值与水分利用效率和土壤氮饱和程度的相互关系,以揭示不同树种水分利用效率、氮饱和程度和碳氮循环速率差异的机理。结果表明:(1)滨海沙地不同树种叶片δ~(13) C变化范围为-31.682‰~-29.323‰,其δ~(13) C大小为:湿地松肯氏相思木麻黄纹荚相思尾巨桉,除尾巨桉外各树种δ~(13) C均表现为新叶老叶;各树种叶片δ~(15)N变化范围为-5.548‰~-2.167‰,其δ~(15)N大小为:肯氏相思纹荚相思木麻黄湿地松尾巨桉,且各树种均表现为新叶老叶。(2)不同树种表层土壤δ~(15)N变化范围为-4.675‰~-2.975‰,表层土壤δ~(15)N大小为:纹荚相思肯氏相思木麻黄尾巨桉湿地松,但不同树种表层土壤C含量无显著差异。(3)滨海沙地湿地松、木麻黄、肯氏相思和纹荚相思的水分利用效率随叶龄增加均呈显著递减趋势;不同树种新叶的水分利用效率变化范围为39.09~76.57μmol·mol~(-1),其大小依次为:湿地松肯氏相思木麻黄纹荚相思尾巨桉;老叶的水分利用效率变化范围为38.56~62.59μmol·mol~(-1),其大小依次为:湿地松木麻黄肯氏相思尾巨桉纹荚相思。(4)不同树种人工林水分利用效率与其新叶水分利用效率呈显著正相关关系,说明林分水分利用效率主要体现在新叶的水分利用效率上,同时林分水分利用效率受林分类型的影响。     

喀斯特槽谷典型植物水分利用效率对隧道建设的响应

隧道工程建设对喀斯特槽谷地区地下水循环系统的破坏,引起地下水位下降,可能影响周围生态环境。依托重庆中梁山喀斯特槽谷,对旱、雨两季隧道影响区与无隧道影响区不同深度土层(0—20 cm、20—40 cm)土壤含水量以及典型植物(常绿乔木、常绿灌木)瞬时水分利用效率(InstantaneousWater Use Efficiency,WUE_inst)进行对比分析,探究植物水分利用效率对隧道建设的响应。结果表明:在不同影响区,主要由蒸发与降水引起土壤含水量在垂直和季节变化上的趋势相似,但隧道影响区土壤含水量在旱、雨两季均显著高于无隧道影响区;不同影响区植物WUE_inst存在明显的季节差异,旱季各植物种WUE_inst显著高于雨季。而旱、雨两季隧道影响区植物叶片WUE_inst显著高于无隧道影响区;此外,植物WUE_inst与土壤含水量相关性分析结果表明,不同影响区植物WUE_inst与土壤含水量均呈显著负相关关系,但相对于无隧道影响区,隧道影响区植物WUE_ins...     

陆生植物水分利用效率

植物水分利用效率（water use efficiency,WUE）作为叶片光合与水分生理过程的耦合因子,不仅是联系植被生态系统碳循环与水循环的重要变量,同时亦已成为揭示陆地植被生态系统对全球变化响应和适应对策的重要手段。稳定同位素技术和涡度相关技术的发展和应用,使植物WUE的概念和测定方法不断得到扩展,研究内容、尺度也不断拓宽。深入认识植物WUE的变化特征及其调控机制,一方面有助于揭示不同植物对生境的生理生态适应机理,另一方面能够促进对陆地生态系统碳-水平衡关系的理解。目前生态系统水平WUE的研究尚处初始阶段。如何建立不同尺度水平WUE间的联系机制,将成为今后该研究领域的关注点。本文在阐述植物WUE概念和测定方法的基础上,综述了植物叶片水平WUE的影响因子、种间差异与水分利用策略,以及WUE与其他叶性状的关系,介绍了当前生态系统水平WUE的研究状况,提出未来的研究趋势。     

陆生植物稳定碳同位素组成与全球变化

分析了大气CO₂浓度、温度、降水和海拔高度等环境因素对陆生植物稳定性碳同位素组分的影响及其作用机理,综述了国内外碳稳定同位素技术在全球变化研究中的进展和应用,如重建大气CO₂浓度变化,揭示温度、降水对树木生长的“滞后效应”和“幼龄效应”,确定不同光合型植物随海拔高度的分布变化,以及通过碳稳定同位素技术揭示不同时间尺度上和不同气候条件下的植物水分利用效率变化及不同生活型植物的水分利用效率差异,并探讨研究中存在的问题及其研究前景.     

Leaf stable carbon isotope composition reflects transpiration efficiency in Zea mays

The increasing demand for food production and predicted climate change scenarios highlight the need for improvements in crop sustainability. The efficient use of water will become increasingly important for rain‐fed agricultural crops even in fertile regions that have historically received ample precipitation. Improvements in water‐use efficiency in Zea mays have been limited, and warrant a renewed effort aided by molecular breeding approaches. Progress has been constrained by the difficulty of measuring water‐use in a field environment. The stable carbon isotope composition (δ¹³C) of the leaf has been proposed as an integrated signature of carbon fixation with a link to stomatal conductance. However, additional factors affecting leaf δ¹³C exist, and a limited number of studies have explored this trait in Z. mays. Here we present an extensive characterization of leaf δ¹³C in Z. mays. Significant variation in leaf δ¹³C exists across diverse lines of Z. mays, which we show to be heritable across several environments. Furthermore, we examine temporal and spatial variation in leaf δ¹³C to determine the optimum sampling time to maximize the use of leaf δ¹³C as a trait. Finally, our results demonstrate the relationship between transpiration and leaf δ¹³C in the field and the greenhouse. Decreasing transpiration and soil moisture are associated with decreasing leaf δ¹³C. Taken together these results outline a strategy for using leaf δ¹³C and reveal its usefulness as a measure of transpiration efficiency under well‐watered conditions rather than a predictor of performance under drought.     

A genetic link between leaf carbon isotope composition and whole‐plant water use efficiency in the C4 grass Setaria

Genetic selection for whole‐plant water use efficiency (yield per transpiration; WUE_plant) in any crop‐breeding programme requires high‐throughput phenotyping of component traits of WUE_plant such as intrinsic water use efficiency (WUE_i; CO₂ assimilation rate per stomatal conductance). Measuring WUE_i by gas exchange measurements is laborious and time consuming and may not reflect an integrated WUE_i over the life of the leaf. Alternatively, leaf carbon stable isotope composition (δ¹³C_leaf) has been suggested as a potential time‐integrated proxy for WUE_i that may provide a tool to screen for WUE_plant. However, a genetic link between δ¹³C_leaf and WUE_plant in a C₄ species has not been well established. Therefore, to determine if there is a genetic relationship in a C₄ plant between δ¹³C_leaf and WUE_plant under well watered and water‐limited growth conditions, a high‐throughput phenotyping facility was used to measure WUE_plant in a recombinant inbred line (RIL) population created between the C₄ grasses Setaria viridis and S. italica. Three quantitative trait loci (QTL) for δ¹³C_leaf were found and co‐localized with transpiration, biomass accumulation, and WUE_plant. Additionally, WUE_plant for each of the δ¹³C_leaf QTL allele classes was negatively correlated with δ¹³C_leaf, as would be predicted when WUE_i influences WUE_plant. These results demonstrate that δ¹³C_leaf is genetically linked to WUE_plant, likely to be through their relationship with WUE_i, and can be used as a high‐throughput proxy to screen for WUE_plant in these C₄ species.     

Photosynthesis, water use efficiency and stable carbon isotope composition are associated with anatomical properties of leaf and xylem in six poplar species

Although fast‐growing Populus species consume a large amount of water for biomass production, there are considerable variations in water use efficiency (WUE) across different poplar species. To compare differences in growth, WUE and anatomical properties of leaf and xylem and to examine the relationship between photosynthesis/WUE and anatomical properties of leaf and xylem, cuttings of six poplar species were grown in a botanical garden. The growth performance, photosynthesis, intrinsic WUE (WUE_i), stable carbon isotope composition (δ¹³C) and anatomical properties of leaf and xylem were analysed in these poplar plants. Significant differences were found in growth, photosynthesis, WUE_i and anatomical properties among the examined species. Populus cathayana was the clone with the fastest growth and the lowest WUE_i/δ¹³C, whereas P. × euramericana had a considerable growth increment and the highest WUE_i/δ¹³C. Among the analysed poplar species, the highest total stomatal density in P. cathayana was correlated with its highest stomatal conductance (g_s) and lowest WUE_i/δ¹³C. Moreover, significant correlations were observed between WUE_i and abaxial stomatal density and stem vessel lumen area. These data suggest that photosynthesis, WUE_i and δ¹³C are associated with leaf and xylem anatomy and there are tradeoffs between growth and WUE_i. It is anticipated that some poplar species, e.g. P. × euramericana, are better candidates for water‐limited regions and others, e.g. P. cathayana, may be better for water‐abundant areas.     

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。     

The physiological basis for genetic variation in water use efficiency and carbon isotope composition in Arabidopsis thaliana

Ecologists and physiologists have documented extensive variation in water use efficiency (WUE) in Arabidopsis thaliana, as well as association of WUE with climatic variation. Here, we demonstrate correlations of whole-plant transpiration efficiency and carbon isotope composition (δ¹³C) among life history classes of A. thaliana. We also use a whole-plant cuvette to examine patterns of co-variation in component traits of WUE and δ¹³C. We find that stomatal conductance (g _s) explains more variation in WUE than does A. Overall, there was a strong genetic correlation between A and g _s, consistent with selection acting on the ratio of these traits. At a more detailed level, genetic variation in A was due to underlying variation in both maximal rate of carboxylation (V _cmax) and maximum electron transport rate (Jmax). We also found strong effects of leaf anatomy, where lines with lower WUE had higher leaf water content (LWC) and specific leaf area (SLA), suggesting a role for mesophyll conductance (g _m) in variation of WUE. We hypothesize that this is due to an effect through g _m, and test this hypothesis using the abi4 mutant. We show that mutants of ABI4 have higher SLA, LWC, and g _m than wild-type, consistent with variation in leaf anatomy causing variation in g _m and δ¹³C. These functional data also add further support to the central, integrative role of ABI4 in simultaneously altering ABA sensitivity, sugar signaling, and CO₂ assimilation. Together our results highlight the need for a more holistic approach in functional studies, both for more accurate annotation of gene function and to understand co-limitations to plant growth and productivity.     

沉水植物稳定碳同位素组成及影响因素分析

青康公路沿线一些河流和湖泊中同种沉水植物－－龙须眼子菜（Potamogeton pectinatus）的稳定碳同位组成（δ^13C）变化在－10.92‰-22.07‰之间,这些龙须眼子菜的δ^13C值大小为盐水湖＞淡水湖＞河流。根据沉水植物的碳同位素分馏模式,对影响龙须眼子菜碳同位素组成的环境因素进行了探讨。结果表明,龙须眼子菜的碳同位素组成受水体环境因素影响。     

Comparison of seasonal variations in water-use efficiency calculated from the carbon isotope composition of tree rings and flux data in a temperate forest

Tree-ring δ(13) C is often interpreted in terms of intrinsic water-use efficiency (WUE) using a carbon isotope discrimination model established at the leaf level. We examined whether intra-ring δ(13) C could be used to assess variations in intrinsic WUE (W(g), the ratio of carbon assimilation and stomatal conductance to water) and variations in ecosystem WUE (W(t) , the ratio of C assimilation and transpiration) at a seasonal scale. Intra-ring δ(13) C was measured in 30- to 60-μm-thick slices in eight oak trees (Quercus petraea). Canopy W(g) was simulated using a physiologically process-based model. High between-tree variability was observed in the seasonal variations of intra-ring δ(13) C. Six trees showed significant positive correlations between W(g) calculated from intra-ring δ(13) C and canopy W(g) averaged over several days during latewood formation. These results suggest that latewood is a seasonal recorder of W(g) trends, with a temporal lag corresponding to the mixing time of sugars in the phloem. These six trees also showed significant negative correlations between photosynthetic discrimination Δ calculated from intra-ring δ(13) C, and ecosystem W(t), during latewood formation. Despite the observed between-tree variability, these results indicate that intra-ring δ(13) C can be used to access seasonal variations in past W(t).     

中国东部南北样带森林优势植物叶片的水分利用效率和氮素利用效率

通过测定中国东部南北样带主要森林生态系统中10种优势植物(兴安落叶松、蒙古栎、水曲柳、紫椴、色木槭、红松、杉木、木荷、马尾松、锥栗)叶片的碳氮含量(Cmass、Nmass)、同位素丰度(δ13C、δ15N)以及光合响应曲线,分析了不同优势植物叶片的水分利用效率和氮素利用效率之间的差异及其相互关系.结果表明： 不同生活型植物叶片的Nmass和δ15N差异显著,表现为阔叶植物>针叶植物,落叶植物>常绿植物;最大光合速率(Pn max)表现为针叶植物>阔叶植物,落叶植物>常绿植物;植物叶片的瞬时水分利用效率(WUEi)和长期水分利用效率(WUE)均表现为阔叶植物>针叶植物,常绿植物>落叶植物;植物叶片的瞬时氮素利用效率(NUEi)和长期氮素利用效率(NUE)则表现出相反的规律,且常绿植物和落叶植物叶片的NUE差异显著;WUEi和WUE之间相关性不显著,而NUEi和NUE之间呈显著正相关.植物叶片的水分利用效率与氮素利用效率显著负相关.两种资源利用效率均受植物生活型的影响,并且存在一定的制约关系.     

Water use efficiency and carbon isotope composition of plants in a cold desert environment

Summary The effects of the availabilities of water and nitrogen on water use efficiency (WUE) of plants were investigated in a sagebrush steppe. The four species studied wereArtemisia tridentata (shrub),Ceratoides lanata (suffrutescent shrub),Elymus lanceolatus (rhizomatous grass), andElymus elymoides (tussock grass). Water and nitrogen levels were manipulated in a two-by-two factorial design resulting in four treatments: control (no additions), added water, added nitrogen, and added water and nitrogen. One instantaneous and two long-term indicators of WUE were used to testa priori predictions of the ranking of WUE among treatments. The short-term indicator was the instantaneous ratio of assimilation to transpiration (A/E). The long-term measures were 1) the slope of the relationship between conductance to water vapor and maximum assimilation and 2) the carbon isotope composition (¹³C) of plant material. Additional water decreased WUE, whereas additional nitrogen increased WUE. For both A/E and ¹³C, the mean for added nitrogen alone was significantly greater than the mean for added water alone, and means for the control and added water and nitrogen fell in between. This ranking of WUE supported the hypothesis that both water and nitrogen limit plant gas exchange in this semiarid environment. The short- and long-term indicators were in agreement, providing evidence in support of theoretical models concerning the water cost of carbon assimilation.