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。     

鬼箭锦鸡儿叶片和土壤碳稳定同位素特征及其影响因素

为探究高海拔地区的植物碳(C)循环过程与其生境的关系,以生长在高山地区的豆科灌木鬼箭锦鸡儿为研究对象,沿着横跨我国东西部山区的样带采集35个样点的鬼箭锦鸡儿叶片和土壤样品,分析了鬼箭锦鸡儿叶片碳稳定同位素组成(δ¹³C)、土壤δ¹³C、叶片和土壤δ¹³C差值(Δδ¹³C)在不同采样点的特征及其与气候因子、叶片和土壤元素的关系。结果表明:鬼箭锦鸡儿叶片δ¹³C的变化范围为-30.9‰～-27.1‰,平均值为-28.4‰,土壤δ¹³C的变化范围为-26.2‰～-23.2‰,平均值为-25.3‰,Δδ¹³C的变化范围为2.0‰～7.7‰,平均值为3.1‰;叶片δ¹³C显著低于土壤δ¹³C,且随着叶片δ¹³C增加,土壤δ¹³C先降低后升高;叶片δ¹³C与生长季均温和叶片C含量呈显著负相关,土壤δ¹³C与相对湿度和最暖月均温呈显著负相关,与土壤碳∶氮(C∶N)呈显著正相关,随土壤C含量的增加土壤δ¹³C先降低后升高,Δδ¹³C与叶片C含量、土壤C含量和土壤C∶N呈显著正相关;气候因子对叶片δ¹³C和Δδ¹³C具有直接影响,同时也通过对叶片和土壤元素的影响,间接导致叶片δ¹³C、土壤δ¹³C和Δδ¹³C的改变。高海拔地区的气候因子、叶片和土壤元素共同影响鬼箭锦鸡儿的C循环过程。     

祁连山两种优势乔木叶片δ13C的海拔响应及其机理

以分布于祁连山北麓中段的两种优势乔木祁连圆柏(Sabina przewalskii)和青海云杉(Picea crassifolia)为研究对象, 分析了高山乔木叶片δ¹³C值对海拔(2 600-3 600 m)、土壤含水量和叶片含水量、叶片碳氮含量的响应及其机理。结果表明, 这两种乔木叶片δ¹³C值均随海拔升高呈增重趋势, 与海拔呈显著正相关关系(p < 0.000 1)。海拔2 600-3 600 m阳坡树种祁连圆柏叶片的δ¹³C值显著高于同海拔梯度阴坡树种青海云杉。祁连圆柏和青海云杉叶片的δ¹³C值均与年平均气温呈显著负相关关系(p < 0.000 1), 与年平均降水量呈显著正相关关系(p < 0.000 1)。祁连圆柏叶片δ¹³C值与土壤含水量(p < 0.000 1)、叶片含水量(p = 0.01)和叶片碳氮比(C/N) (p < 0.000 1)呈显著正相关关系, 与叶片全氮呈显著负相关关系(p < 0.000 1)。而青海云杉叶片δ¹³C值与土壤含水量、叶片全氮、叶片碳氮比和叶片含水量不相关。说明海拔变化引起的水热条件的改变, 尤其是温度变化对高山乔木叶片碳同位素分馏起主要作用, 但各个因子综合对高山植物叶片碳同位素分馏的作用机制可能比较复杂, 需进一步深入研究。     

沙棘属植物叶片碳稳定同位素含量与气候的关系

本试验以131个沙棘属植物种群为研究对象,通过测定其叶片碳稳定同位素(δ¹³C)值,分析了碳稳定同位素特征与环境因子之间的关系。结果表明: 沙棘属植物叶片的δ¹³C值介于-24.65‰～-29.11‰,平均值为-26.97‰,属于C3植物,叶片δ¹³C值变异系数为种内大于种间,表明环境因子是影响沙棘属植物叶片δ¹³C含量变化的主导因素。沙棘属植物叶片的δ¹³C值与经纬度的变化无显著相关,与海拔呈显著负相关。通过建立回归方程: δ¹³C(‰)=0.118VAP-0.007GST-0.000028RDA-20.721(R²=0.212,P<0.0001),说明影响沙棘属叶片δ¹³C值最主要的因素是水蒸气压(VAP)、生长季温度(GST)和太阳辐射(RDA)。研究结果可为沙棘属植物对全球气候变化的响应提供理论依据。     

不同基因型冬小麦旗叶的稳定碳同位素比值及其与产量和水分利用效率的关系

以我国北方12个冬小麦(Triticum aestivum)品种(系)和美国德克萨斯州3个冬小麦品种(系)为供试材料, 在甘肃陇东黄土高原旱作和拔节期有限补灌条件下, 比较研究了不同基因型冬小麦之间产量、水分利用效率(WUE)和灌浆期旗叶稳定碳同位素比值(δ¹³C)的差异, 以及δ¹³C值与产量和WUE的关系。旨在通过分析δ¹³C值与产量和WUE的关系, 明确δ¹³C值在评价植物WUE方面的可靠性, 为抗旱节水品种的筛选提供理论依据。结果表明: 不论旱作还是有限补灌, 不同基因型冬小麦之间产量、WUE、旗叶δ¹³C值存在显著差异, 随着灌浆过程的进行, 旗叶δ¹³C值呈缓慢增大的趋势, 而且旗叶δ¹³C值旱作高于有限补灌。不论旱作还是补灌条件, 旗叶δ¹³C值在4个测定时期的平均值与籽粒产量、WUE呈显著正相关关系(R²= 0.527 3-0.691 3)。小麦拔节期补灌100 mm水分后, 不同基因型小麦表现出明显的水分超补偿效应。说明冬小麦灌浆期旗叶δ¹³C值在旱作条件下和在补灌条件下均可较好地评价WUE, 可将冬小麦灌浆期旗叶δ¹³C值作为筛选高效用水品种的参考指标之一。     

不同生境间红树科植物水分利用效率的比较研究

通过测定采自4个地区(海南、厦门、北海和西双版纳)的红树科6个属共9种植物,包括竹节树(Carallia brachiata)、锯叶竹节树(C. diphopetala)、山红树(Pellacalyx yunnanensis)、红树(Rhizophora apiculata)、红海榄(R. stylosa)、海莲(Bruguiera sexangula)、木榄(B. gymnorhiza)、秋茄(Kandelia candel)和角果木(Ceriops tagal)的叶片碳同位素比值(δ¹³C),比较了不同地区分布的红树科植物(尤其是内陆生长的和沿海生长的红树科植物之间)、同一地区分布不同种红树科植物间以及不同季节红树科植物δ¹³C值及其所反映的胞间CO₂浓度和水分利用效率的差异。研究结果表明,红树科植物叶片的δ¹³C变化在-32‰~-26‰之间,大部分种类在两个生长季之间(春季和秋季)没有明显的差异,而内陆和沿海分布的红树科植物有着显著不同的δ¹³C值,以海水中生长的红树科植物δ¹³C值较高。此外,在海水中生长的红树科植物以北海地区分布的为最高,而在厦门和海南之间则较少有显著性的差异。从所取得的结果来看,植物δ¹³C值之间的差异可能有遗传学的基础,但环境的影响也起很大的作用。     

毛白杨杂种无性系叶片δ13C差异与气体交换参数

在苗木生长的不同时期对13个毛白杨(Populus tomentosa)杂种无性系叶片碳同位素δ¹³C和气体交换参数(净光合速率P_n、蒸腾速率T_r、瞬时水分利用效率WUE_i、气孔导度G_s和胞间CO₂浓度C_i)的差异进行研究, 分析不同无性系间δ¹³C与气体交换参数的相互关系, 目的在于探求δ¹³C在筛选高光合及高水分利用效率毛白杨杂种无性系中的应用价值。结果表明: 不同生长时期和不同无性系间δ¹³C、T_r、WUE_i、G_s和C_i的差异均显著, δ¹³C和WUE_i表现为9月>7月, T_r、G_s和C_i表现为7月>9月, P_n在不同生长时期差异不显著。季节变化是引起毛白杨杂种无性系叶片δ¹³C差异的主要原因。同一时期, 无性系间δ¹³C和WUE_i表现出较好的一致性, 即WUE_i较高的无性系30、42、46、83、BL2和BL5, 其δ¹³C值也较高, WUE_i较低的无性系B331和TG34, 其δ¹³C值也较低, 且不同时期(7月和9月) δ¹³C和WUE_i呈较强的正相关, 相关系数r分别为0.739 0和0.545 8, 高δ¹³C可以作为筛选高WUE_i毛白杨的有效指标, 且在苗木生长旺盛时期选育能得到更为可靠的结果。对毛白杨而言, 高WUE_i的无性系, 一般具有适中或较低的G_s和C_i, 但不一定具有很高的P_n, 气孔调节使得毛白杨在不影响光合作用的同时保持较高的WUE。     

全球木本植物叶片硅钙生态化学计量学特征

收集全球803种木本植物叶片硅（Si）、钙（Ca）数据,研究不同木本植物生活型（常绿植物以及落叶植物、针叶植物以及阔叶植物）叶片Si、Ca元素的化学计量学特征及其与纬度、气候因子（年平均温度,年平均降水量）间的关系。结果表明：（1）全球尺度上木本植物叶片Si、Ca含量存在较大变异性,且含量均低于中国境内的研究结果;（2）不同生活型树种间存在差异,针叶树叶片Si含量及Ca/Si显著高于阔叶树,落叶树叶片Si、Ca含量及Ca/Si均显著高于常绿树种;（3）随着纬度升高、年平均温度及年平均降水量的下降,全球尺度木本植物叶片Si、Ca含量显著升高,而Ca/Si显著下降;（4）不同生活型木本植物对气候因子的响应不同,除针叶及落叶树种的Ca含量外,其余各生活型树种Si、Ca含量与纬度及气候因子显著相关,随着纬度升高而升高,随年平均温度及年平均降水量的升高而降低,且随着年平均温度的降低,常绿及阔叶树种叶片Si含量下降速度显著高于落叶及针叶树种。研究结果能够为全球尺度生态化学计量学模型的发展提供数据基础,有助于更好地理解和模拟区域乃至全球尺度上纬度和气候因子对植物叶片Si、Ca含量的影响。     

天津沼泽湿地芦苇叶片的碳稳定同位素比值分布特征及其环境影响因素

天津芦苇(Phragmites australis)沼泽具有重要的生态功能。目前天津地区水体咸化、氮污染和水资源短缺问题严重, 显著影响了芦苇湿地的植物生理生态过程。植物稳定碳同位素组成(碳稳定同位素比值(δ¹³C))能够记录与植物生长过程相关联的环境变化信息, 反映植物对环境变化的生理生态适应特性。该研究调查了天津七里海、北大港和大黄堡湿地芦苇叶片的δ¹³C分布特征, 探讨了影响该地区叶片δ¹³C值变化的主要因素。研究表明: 1)天津芦苇湿地植物叶片δ¹³C的变化范围在-26.3‰ - -23.6 ‰之间, 平均值为-25.8‰; 2)芦苇叶片δ¹³C与底泥相对含水量呈显著负相关关系, 与底泥有效氮和全氮含量呈显著正相关关系, 而与底泥盐度和磷含量没有显著相关关系; 水分条件和底泥氮营养状况是影响叶片的δ¹³C值变化的主要因素; 3)淹水条件下, 芦苇叶片δ¹³C与叶片质量氮含量呈显著正相关关系, 与叶碳氮比呈显著负相关关系, 8月份七里海湿地干涸打破了此相关关系。当前环境压力下, 天津沼泽湿地干涸极大地改变了芦苇的氮、水平衡和植物对水、氮资源的利用策略, 而湿地干涸对该过程的影响要高于盐度和氮负荷增加。     

A review of the calibration methods for measuring the carbon and oxygen isotopes in CO2 based on isotope ratio infrared spectroscopy

稳定同位素红外光谱(IRIS)技术克服了传统的大气CO₂气瓶采样-同位素质谱(IRMS)技术时间分辨率低且耗时费力的缺点, 可以实现高时间分辨率和高精度的大气CO₂碳同位素组成(δ ¹³C)和氧同位素组成(δ ¹⁸O)的原位连续测定。基于IRIS技术测量CO₂ δ ¹³C和δ ¹⁸O的误差来源主要包括δ ¹³C和δ ¹⁸O测量值对CO₂浓度变化的非线性响应(浓度依赖性)以及对环境条件变化的敏感性导致的漂移(时间漂移)。如何有效地校正浓度依赖性和时间漂移导致的误差是IRIS仪器应用的前提。该综述阐述了δ ¹³C和δ ¹⁸O测量值的浓度依赖性产生的理论基础, 回顾了浓度依赖性的理论校正和经验方程校正方法和应用; 回顾了时间漂移的校正原理、方法和应用; 概述了数据溯源至国际标准的原理、方法与应用现状。结合实际情况推荐利用3个或3个以上已知CO₂浓度和δ ¹³C、δ ¹⁸O真值的CO₂标准气体涵盖待测气体CO₂浓度的浓度依赖性校正, 设置适当的校正频率校正时间漂移并进行数据溯源。指出应该加强不同仪器和校正方法的比对研究; 采用IRIS技术测定CH₄、N₂O和H₂O同位素组成也可以采取类似的校正方法。     

Variation in foliar δ13C in Hawaiian Metrosideros polymorpha: a case of internal resistance?

Summary Sun leaves of Metrosideros polymorpha were collected in 51 sites on 9 lava flows that represented gradients of elevation, precipitation, substrate age, and substrate texture on Mauna Loa volcano, Hawai'i. Leaf mass per unit leaf area increased with increasing elevation on all flows, while foliar nitrogen concentration decreased with increasing elevation and increased with increasing substrate age. Foliar ¹³C became less negative with increasing elevation on the wet east-side lava flows, but not the dry northwest-side flows; it did not reflect patterns of precipitation or presumed water availability. ¹³C was very strongly correlated with leaf mass per area across all of the sites. Limited gas-exchange information suggested that calculated c_i/c_a did not decrease with elevation in association with less-negative ¹³C, and photosynthesis per unit of nitrogen was significantly reduced in high-elevation plants. These results are consistent with a substantial internal resistance to CO₂ diffusion in the thick Metrosideros polymorpha leaves in high elevation sites.Published as a contribution to the International Union for Biological Sciences Tropical Mountain Ecosystems Program, and as Carnegie Institution of Washington Department of Plant Biology Publication Number 1065     

长期垄作稻田腐殖质稳定碳同位素丰度(δ13C)分布特征

研究了四川盆地丘陵区连续16年垄（宽垄）作稻田土壤稳定碳库腐殖质组分的稳定碳同位素(δ¹³C)分布特征.结果表明： 稻田土壤有机碳含量为宽垄作＞垄作＞水旱轮作.腐殖质碳以胡敏素为主,占土壤碳含量的21%～30%,提取碳以胡敏酸为主,分别占土壤有机碳和腐殖质的17%～21%和38%～65%.土壤有机碳的δ¹³C值介于-27.9‰～-25.6‰,20～40 cm和0～5 cm土壤有机碳δ¹³C值之差约为1.9‰.土壤胡敏酸δ¹³C值比土壤有机碳低1‰～2‰,更接近于油菜和水稻秸秆及根系的δ¹³C值.土壤富里酸δ¹³C值分别较土壤有机碳和胡敏酸高2‰和4‰.耕作层和犁底层胡敏素δ¹³C值分别介于-23.7‰～-24.9‰和-22.6‰～-24.2‰,δ¹³C值的变化反映了耕层中腐殖质的新老混合现象.各有机组分δ¹³C值递减顺序为：胡敏素＞富里酸＞土壤有机碳＞稻草（油菜）残体＞胡敏酸.长期水稻种植有利于增加土壤有机碳含量,同时,耕作方式影响土壤腐殖质δ¹³C在耕作层和犁底层中的分布格局.     

Measurement of rhizosphere respiration and organic matter decomposition using natural 13C

Due to the limitations in methodology it has been a difficult task to measure rhizosphere respiration and original soil carbon decomposition under the influence of living roots. ¹⁴C-labeling has been widely used for this purpose in spite of numerous problems associated with the labeling method. In this paper, a natural ¹³C method was used to measure rhizosphere respiration and original soil carbon decomposition in a short-term growth chamber experiment. The main objective of the experiment was to validate a key assumption of this method: the ¹³C value of the roots represents the ¹³C value of the rhizosphere respired CO₂. Results from plants grown in inoculated carbon-free medium indicated that this assumption was valid. This natural ¹³C method was demonstrated to be advantageous for studying rhizosphere respiration and the effects of living roots on original soil carbon decomposition.     

Climatic adaptability of populations ofDiplotaxis erucoides D.C. (Brassicaceae) from Sicily,based on leaf morphology,leaf anatomy and δ13 C studies

The morphological and anatomical variability ofDiplotaxis erucoides populations from Sicily was investigated. Populations growing during the summer months exhibit distinct xeromorphic features. Leaf area is strongly reduced and leaf thickness is increased when compared with winter populations. Cell size, as well as cell arrangement and mesophyll cell surface area differ significantly between summer and winter populations. Leaf thickness is almost three times higher in summer populations andA (cell)/A, i.e. the mesophyll cell surface area per unit leaf area changes from about 16 for winter populations to almost 52 for summer populations. These differences are partly due to differences in intercellular volume and partly due to alterations in mesophyll cell sizes. The organic materal of the summer populations exhibits ¹³C values in the order of –27%. to –28%., while the corresponding values for the winter populations are in the order of –31%. to –33%.. Analysis ofD. erucoides populations from the transition period revealed intermediate ¹³C values. Anatomical variations such as reductions or increases ofA (cells)/A and changes of intercellular volume correlate with the corresponding ¹³C data. The ¹³C data are discussed in conjunction with the differences in leaf anatomy.     

Foliar δ<Superscript>13</Superscript>C values of nine dominant species in the Loess Plateau of China

The foliar stable carbon isotope compositions (δ¹³C) of nine dominant species in seven sites, Yangling, Yongshou, Tongchuan, Fuxian, Ansai, Mizhi, and Shenmu, standing from the south to the north in the Loess Plateau of China were studied. The results showed that foliar δ¹³C values ranged from −22.61 to −30.73 ‰ with an average of −27.23 ‰ in 141 C₃ plant samples collected from the Loess Plateau. Foliar δ¹³C values varied significantly (p<0.001) among the nine C₃ species, which were Pinus tabulaeformis Carr., Robinia pseudoacacia L., Zizyphus jujuba Mill. var. spinosus Hu., Rubus parvifolius L., Hippophae rhamnoides L., Caragana korshinskii Kom., Lespedeza davurica (Laxm.) Schindl., Artemisia sacrorum Ledeb. var. incana Mattf., and Agropyron cristatum Gaertn. Comparatively, R. pseudoacacia, H. rhamnoides, and C. korshinskii had much higher δ¹³C values than the other six species, while A. sacrorum had the lowest δ¹³C value. There was no significant difference in foliar δ¹³C value among five species, P. tabulaeformis, Z. jujuba, R. parvifolius, L. davurica, and A. cristatum. Considering the life forms categorized from nine C₃ species, trees and shrubs had significantly higher δ¹³C values than herbs (p<0.001). The deciduous tree R. pseudoacacia had much higher δ¹³C value than the evergreen tree P. tabulaeformis (p<0.01). Among the four shrubs, foliar δ¹³C values in H. rhamnoides and C. korshinskii were markedly higher (p<0.01) than those in Z. jujuba and R. parvifolius. Among the three herbs, L. davurica and A. cristatum had significantly higher δ¹³C values than A. sacrorum (p<0.01). Leguminous species such as R. pseudoacacia, C. korshinskii, and L. davurica as well as a non-leguminous species with nitrogen-fixation capacity, H. rhamnoides, had higher δ¹³C values than other non-leguminous species with same life-form. The mean δ¹³C value increased by about 7 % from Yangling in the south to Shenmu in the north as climatic drought increased, and foliar δ¹³C values differed much (p<0.001) among the seven sites. For nine species in the Loess Plateau, foliar δ¹³C values were significantly and negatively (p<0.001) correlated with the mean annual precipitation, moreover, an increase of 100 mm in annual precipitation would result in a decrease of 1.2 ‰ in δ¹³C value.     

Foliar δ13C within a temperate deciduous forest: spatial,temporal, and species sources of variation

Summary Foliar ¹³C-abundance (¹³C) was analyzed in the dominant trees of a temperate deciduous forest in east Tennessee (Walker Branch Watershed) to investigate the variation in foliar ¹³C as a function of time (within-year and between years), space (canopy height, watershed topography and habitat) and species (deciduous and coniferous taxa). Various hypotheses were tested by analyzing (i) samples collected from the field during the growing season and (ii) foliar tissues maintained in an archived collection. The ¹³C-value for leaves from the tops of trees was 2 to 3%. more positive than for leaves sampled at lower heights in the canopy. Quercus prinus leaves sampled just prior to autumn leaf fall had significantly more negative ¹³C-values than those sampled during midsummer. On the more xeric ridges, needles of Pinus spp. had more positive ¹³C-values than leaves from deciduous species. Foliar ¹³C-values differed significantly as a function of topography. Deciduous leaves from xeric sites (ridges and slopes) had more positive ¹³C-values than those from mesic (riparian and cove) environments. On the more xeric sites, foliar ¹³C was significantly more positive in 1988 (a dry year) relative to that in 1989 (a year with above-normal precipitation). In contrast, leaf ¹³C in trees from mesic valley bottoms did not differ significantly among years with disparate precipitation. Patterns in foliar ¹³C indicated a higher ratio of net CO₂ assimilation to transpiration (A/E) for trees in more xeric versus mesic habitats, and for trees in xeric habitats during years of drought versus years of normal precipitation. However, A/E (units of mmol CO₂ fixed/mol H₂O transpired) calculated on the basis of ¹³C-values for leaves from the more xeric sites was higher in a wet year (6.6±1.2) versus a dry year (3.4±0.4). This difference was attributed to higher transpiration (and therefore lower A/E) in the year with lower relative humidity and higher average daily temperature. The calculated A/E values for the forest in 1988–89, based on ¹³C, were within ±55% of estimates made over a 17 day period at this site in 1984 using micrometeorological methods.     

Association between tree-ring and needle δ<Superscript>13</Superscript>C and leaf gas exchange in <Emphasis Type="Italic">Pinus halepensis</Emphasis> under semi-arid conditions

Associations between δ¹³C values and leaf gas exchanges and tree-ring or needle growth, used in ecophysiological compositions, can be complex depending on the relative timing of CO₂ uptake and subsequent redistribution and allocation of carbon to needle and stem components. For palaeoenvironmental and dendroecological studies it is often interpreted in terms of a simple model of δ¹³C fractionation in C₃ plants. However, in spite of potential complicating factors, few studies have actually examined these relationships in mature trees over inter- and intra-annual time-scales. Here, we present results from a 4 years study that investigated the links between variations in leaf gas-exchange properties, growth, and dated δ¹³C values along the needles and across tree rings of Aleppo pine trees growing in a semi-arid region under natural conditions or with supplemental summer irrigation. Sub-sections of tissue across annual rings and along needles, for which time of formation was resolved from growth rate analyses, showed rapid growth and δ¹³C responses to changing environmental conditions. Seasonal cycles of growth and δ¹³C (up to ~4‰) significantly correlated (P<0.01) with photosynthetically active radiation, vapour pressure deficit, air temperature, and soil water content. The irrigation significantly increased leaf net assimilation, stomatal conductance and needle and tree-ring growth rate, and markedly decreased needle and tree-ring δ¹³C values and its sensitivity to environmental parameters. The δ¹³C estimates derived from gas-exchange parameters, and weighted by assimilation, compared closely with seasonal and inter-annual δ¹³C values of needle- and tree-ring tissue. Higher stomatal conductances of the irrigated trees (0.22 vs. 0.08 mol m⁻² s⁻¹ on average) corresponded with ~2.0‰ lower average δ¹³C values, both measured and derived. Derived and measured δ¹³C values also indicated that needle growth, which occurs throughout the stressful summer was supported by carbon from concurrent, low rate assimilation. For Aleppo pine under semi-arid and irrigated conditions, the δ¹³C of tree-ring and needle material proved, in general, to be a reasonable indicator of integrated leaf gas-exchange properties.     

Measurements of abundances of 15N and 13C as tools in retrospective studies of N balances and water stress in forests: A discussion of preliminary results

Preliminary attempts to make retrospective studies of N balances and water stress in forest fertilization experiments by analyzing changes in the abundances of ¹⁵N and ¹³C, respectively, are discussed. Most evidence is from the Swedish Forest Optimum Nutrition Experiments, which have been running for two decades. Annual additions of N have been given either alone or in combination with other elements, notably P and K, every third year. Processes leading to loss of N, e.g. volatilization of ammonia, nitrification followed by leaching or denitrification, and denitrification alone, discriminate against the heavy isotope ¹⁵N. A correlation was found between fractional losses of added N and the change in ¹⁵N () during 19 years in current needles in a Scots pine forest, irrespective of source of N. Isotope effects were larger on urea than on ammonium nitrate plots (2 as compared to 9 ¹⁵N ()) because of ammonia volatilization and higher rates of nitrification. They developed gradually over time, which opens possibilities to analyse the development of N saturation. However, the analysis may be confounded by shifts in ¹⁵N abundance of fertilizer N. In another trial, N isotope effects could be seen in both plants and soils 10 years after the last fertilization; they were smaller in soils because of a large pretreatment memory effect, but we expect them to persist there for decades.The enzyme RuBisCo discriminates strongly against the heavy isotope ¹³C during photosynthesis, but this effect becomes less expressed as stomata close because of water stress. The supply of N may also affect the ¹³C () via effects on rates of photosynthesis, and the source of N may have an influence directly via non-RubisCo carboxylations, and indirectly via effects on water use efficiency. In a trial with Norway spruce, the effect of N fertilization on the ¹³C () of current needles was strongly correlated with production and weakly so with foliar biomass a dry year, but not a wet year. This suggested that these variations are primarily related to induced differences in the balance between supply and demand for water. Hence, studies of {au13}C abundance can disentangle the role of water as such from its effects on mineralization of N and flow of N.