高山植物叶片δ 13C的海拔响应及其机理

植物13C的分辨研究已成为植物生态学和全球碳循环研究的核心问题之一.植物13C的分辨是环境和生物因子共同作用的综合结果,海拔梯度变化不仅可以造成植物生存环境的变化,而且还可以造成植物形态和生理特征的变化,因此,高山植物13C分辨随海拔的变化为深入揭示植物13C分辨的环境和生物因子的作用机理提供了非常理想的研究条件.在简单介绍植物13C分辨基本理论的基础上,对目前国际上高山植物13C分辨的海拔响应研究进行了述评.重点介绍了随海拔变化的大气13C组成、温度、气压、水分等环境因子和植物叶片的气孔导度、羧化效率、氮含量和叶肉细胞导度等生物因子对高山C3植物13C分辨的影响,指出高山植物13C分辨的海拔响应机理仍存在一些不确定性,为国内相关研究的开展提供了一定参考.     

青藏高原东部典型高山植物叶片δ13C的季节变化

通过对青藏高原高寒草甸生态系统28种高山植物叶片不同月份稳定碳同位素组成的测定,研究植物δ 13C值在不同季节变化及其与环境之间的关系,试图找出影响δ 13C值变化的关键环境因子.结果表明植物δ13C值在不同月份间有显著性差异(P<0.01),生长初期(6月)δ13C值明显高于生长末期(8月).植物的δ13C值变化主要是由于温度和降水引起的,随温度和降雨量降低而偏重.另外,不同生长期植物叶片的成熟度可能对植物δ13C的变化有一定的贡献.不同种植物稳定性碳同位素值变化差别很大,反应了不同植物对环境变化的不同响应.     

δ13C在植物生态学研究中的应用

稳定碳同位素技术已成为研究植物与环境之间关系最有效的方法之一。由于植物羧化效率的不同、12 C和13 C在植物体内迁移速率以及外界环境的不同,不同植物体内稳定性碳同位素比率(δ13 C值)有一定的差异。该文概述了稳定碳同位素的基本理论,并从气孔导度、叶肉细胞导度、叶片羧化效率分析了δ13 C变化的生物学机理;对近年来国内外有关不同环境因子对植物δ13 C值的影响、δ13 C值在群落及生态系统水平(以功能群、群落冠层及树轮为重点)、以及δ13 C值在碳循环中的应用研究进展进行综述,为以后稳定碳同位素研究提供参考。     

基于树轮δ13C的北京山区侧柏水分利用效率

树木水分利用效率特征存在地域及树种差异,北京山区作为华北土石山区典型森林生态系统,其主要树种侧柏水分利用效率(WUE)长期变化的研究未见报道.本研究通过测定侧柏树轮稳定碳同位素值(δ¹³C),计算年均水分利用效率(WUE_i),分析其长期变化趋势及对环境条件变化的响应,并结合树轮宽度,探究侧柏WUE_i与净固碳量的关系.结果表明: 1918—2013年期间,北京山区年均气温呈逐渐升高趋势,年降水量波动较大.在此期间,侧柏树轮δ¹³C逐渐减小,WUE_i呈逐渐增大趋势.侧柏WUE_i对气温变化的响应最为敏感,呈显著正相关关系,且WUE_i对气温升高的响应敏感性大于气温降低.侧柏WUE_i与年降水量的相关性不明显,该地区降水量不是影响侧柏WUE_i的主要因素.侧柏去趋势树轮宽度值呈现先上升后下降趋势,近20年下降趋势明显.结合WUE_i与环境因子的相关性分析,认为气温升高导致气孔导度(g_s)降低,蒸腾量减少,同时增加了呼吸损耗,结果导致WUE_i升高,但侧柏净固碳量减少,生长减缓.     

栓皮栎叶片δ13C和δ15N的纬向趋势及其影响因子

以东亚广布种栓皮栎为例,通过对南北样带(26°-40° N)上7个群体的调查取样,并结合2个生活史阶段,探讨栓皮栎碳氮同位素比值随纬度环境的变异规律.结果表明: 在纬向梯度上,栓皮栎叶片δ¹³C和δ¹⁵N随纬度的增加分别呈现非线性的增加和下降趋势,且成年树叶片δ¹³C和δ¹⁵N均显著高于幼树;同时,树龄和纬度对叶片δ¹⁵N和δ¹³C均无显著交互作用,表明栓皮栎幼树和成年树纬向环境变化的响应较为一致.随机森林模型结果显示,栓皮栎叶片δ¹⁵N主要受土壤养分,如土壤有机质、磷和氮含量的影响,而叶片δ¹³C主要受水分因子,如空气相对湿度、降水量等的影响.     

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。     

巴郎山刺叶高山栎叶片δ13C对海拔高度的响应

在卧龙自然保护区,按海拔梯度选择了三个刺叶高山栎分布地点(2600m、3050m和3500m),对各研究地点刺叶高山栎进行了叶片d¹³C、光合、CO2扩散导度、养分以及形态等参数的测量,以期揭示刺叶高山栎叶片d¹³C随着海拔的响应规律及原因。结果表明：首先,随着海拔的升高,目标树种叶片d¹³C随之增加,且海拔每升高1000m,d¹³C增加2.0‰;其次,随着海拔升高,大气压强降低,因而导致的叶片扩散导度(包括气孔导度(gs)和叶肉细胞导度(gm))的降低是叶片d¹³C随着海拔升高而升高的主要原因;同时,随着海拔升高而增加的叶片厚度(LMA)也在一定程度上促进了这种响应趋势;最后,叶片N含量随着海拔升高而更多被分配给非光合系统的趋势促进了羧化效率及光合速率的降低,进而对刺叶高山栎叶片d¹³C海拔响应起到了一定的抑制作用,但程度不大;此外,光合氮利用效率(PNUE)在反映植物叶片的氮含量与其吸收和固定大气CO2能力之间的关系方面及对d¹³C的指示作用方面较单位面积氮含量(Narea)和单位重量氮含量(Nmass)更鲜明和准确。     

高寒沙地不同恢复年限沙棘叶片δ13C和δ15N变化特征及其影响因素

人工植被建植是青海湖流域沙地生态恢复的有效措施,揭示典型固沙植物的水分和养分利用效率,并分析影响其时间和空间变化的因素,有利于更好地理解固沙植被对高寒干旱环境有限资源的利用策略,为促进脆弱沙区的植被恢复提供有效的理论依据和实践指导。以青海湖东沙地不同恢复年限(1987年、2008年和2015年)栽植的沙棘为研究对象,利用稳定碳氮同位素技术测定植物叶片的δ¹³C、δ¹⁵N、单位质量叶的碳(C)含量(%)、氮(N)含量(%)并计算叶的碳氮比(C/N),基于土壤含水量、叶片含水量及局地气象因子等要素,探究δ¹³C和δ¹⁵N在季节和空间尺度的变化特征及其对局地土壤和气候环境的响应。结果表明:(1)在时间尺度上,沙棘叶片的δ¹³C与δ¹⁵N在生长季呈现出先减小后增大再减小的趋势,C含量呈先升高后降低再升高的趋势,N含量呈逐渐降低的趋势,而C/N则呈现逐渐升高的趋势。(2)在空间尺度上,幼龄林的δ¹³C最高而老龄林最低,δ¹⁵N值呈现相反的趋势,沙棘叶片的C含量、N含量及其C/N均没有表现出明显的差异。(3)δ¹³C值在不同恢复年限沙棘中受到不同因子的影响,而δ¹⁵N值与平均相对湿度(MRH)之间具有显著的负相关性(P < 0.05)。沙棘δ¹³C值的季节变化是气象因子综合作用的结果,而δ¹⁵N值的季节变化是局地气象因子与叶片养分含量综合作用的结果。随着固沙年限的增加,沙棘的δ¹³C值逐渐降低,而δ¹⁵N值却在逐渐增加,固沙植被从幼龄向老龄演替进程中植物的水分利用效率(WUE)与氮素利用效率(NUE)之间具有权衡关系。     

内蒙古草原常见植物叶片δ13C和δ15N对环境因子的响应

在中国东北样带沿线的内蒙古草原地区采集了一些常见植物的叶片样品,并测定其δ¹³C和δ¹⁵N值,分析了其统计学特征以及对环境因子(年平均降雨量和温度)的响应模式。发现东北样带草原区同时存在C₃和C₄两种不同光合途径的植物,但是C₃植物占主导地位,C₄植物数量有限。C₃植物叶片δ¹³C随着年平均降雨量和年平均温度的升高而显著降低,反映了此区域C₃植物δ¹³C受控于降水量和温度。C₄植物的叶片δ¹³C值随着降雨量的增多而有轻微升高的趋势,但是C₄植物的叶片δ¹³C值对年平均温度的响应不敏感。不论对C₃植物还是C₄植物而言,叶片δ¹⁵N都随降雨量增加而显著降低,即干旱区的植物叶片δ¹⁵N大于湿润地区,这说明降水是影响植物叶片δ¹⁵N的一个重要因素。然而两者叶片δ¹⁵N对温度的响应不敏感。     

水肥耦合效应对楸树苗木叶片δ13C的影响

以楸树(Catalpa bungei)无性系004-1苗木为研究对象,采用三因素五水平二次回归通用旋转组合设计进行盆栽试验,通过测定6、7和8月份20个处理楸树苗木叶片碳同位素组成δ~(13)C、总氮含量和总磷含量,建立数学模型,分析土壤水分、施氮量和施磷量的主因子效应、单因素效应及其耦合效应对叶片δ~(13)C的影响,以及叶片碳同位素组成δ~(13)C与其总氮含量和总磷含量的关系。结果表明:(1)楸树苗木叶片δ~(13)C存在明显的月动态变化特征,总体上呈现出6月份高于7、8月份的趋势,20个处理苗木叶片δ~(13)C平均值(6~8月)变化范围介于-29.12‰~-26.60‰之间;(2)土壤水分对叶片δ~(13)C有显著负效应,施氮量对叶片δ~(13)C有显著正效应,并且施氮量主效应大于土壤水分主效应,但施磷量对叶片δ~(13)C无显著主效应;单因素效应分析表明,楸树苗木叶片δ~(13)C随土壤水分或施氮量的增加均呈先增加后减小的\"抛物线型\"关系;土壤水分×施N量对楸树苗木叶片δ~(13)C有显著耦合负效应,土壤水分和施氮量耦合效应图表明,叶片δ~(13)C随土壤水分降低、施氮量增加而逐渐增加;(3)6月、7月和8月份楸树苗木叶片δ~(13)C与其叶片总氮含量均呈显著正相关关系(P0.05;n=100),但叶片δ~(13)C与总磷含量无显著相关关系(P0.05;n=100)。研究认为,楸树苗木叶片碳同位素组成δ~(13)C随生长期表现出动态变化特征,它受到土壤水分、水氮耦合的显著负向影响,以及施氮量的显著正向影响,可以通过合理水肥配施措施大幅提高楸树苗木δ~(13)C。     

CO2 transfer conductance, leaf structure and carbon isotope composition of Polygonum cuspidatum leaves from low and high altitudes

Anatomy and some physiological characteristics of the leaves in Polygonum cuspidatum Sieb. et Zucc., a dioecious clonal herb, were compared between two populations, one from a lowland in Shizuoka City (10 m above sea level), and another from a highland on Mt. Fuji (2500 m above sea level). Leaf mass per area (LMA) of the highland plants was about twice that of the lowland plants. The greater leaf thickness, thicker mesophyll cell walls and higher mesophyll cell density in the highland leaves contributed to the larger LMA. Although mesophyll area exposed to intercellular airspaces was greater in the highland leaves than in the lowland leaves by 30%, the surface area of chloroplasts facing intercellular airspaces was similar between these leaves. CO₂ transfer conductance inside the leaf (g_i) of the highland leaves (0·75 μmol m^?2 s^?1 Pa^?1) is the lowest recorded for herbaceous plants and was only 40% of that in the lowland leaves. On the other hand, the difference in stomatal conductance was small. δ¹³C values in the leaf dry matter were greater in the highland leaves by 4‰. These data and the estimation of CO₂ partial pressures in the intercellular air spaces and in the chloroplast suggested that the greater dry matter δ¹³C in the highland leaves, indicative of lower long‐term ratio of the chloroplast stroma to the ambient CO₂ partial pressures, would be mainly attributed to their lower g_i.     

Partitioning net ecosystem carbon exchange and the carbon isotopic disequilibrium in a subalpine forest

We investigate the utility of an improved isotopic method to partition the net ecosystem exchange of CO₂ (F) into net photosynthesis (F_A) and nonfoliar respiration (F_R). Measurements of F and the carbon isotopic content in air at a high‐elevation coniferous forest (the Niwot Ridge AmeriFlux site) were used to partition F into F_A and F_R. Isotopically partitioned fluxes were then compared with an independent flux partitioning method that estimated gross photosynthesis (GEE) and total ecosystem respiration (TER) based on statistical regressions of night‐time F and air temperature. We compared the estimates of F_A and F_R with expected canopy physiological relationships with light (photosynthetically active radiation) and air temperature. Estimates of F_A and GEE were dependent on light as expected, and TER, but not F_R, exhibited the expected dependence on temperature. Estimates of the isotopic disequilibrium D , or the difference between the isotopic signatures of net photosynthesis (δ_A, mean value ?24.6‰) and ecosystem respiration (δ_R, mean value ?25.1‰) were generally positive (δ_A>δ_R). The sign of D observed here is inconsistent with many other studies. The key parameters of the improved isotopic flux partitioning method presented here are ecosystem scale mesophyll conductance (g_m) and maximal vegetative stomatal conductance (g_cmax). The sensitivity analyses of F_A, F_R, and D to g_cmax indicated a critical value of g_cmax (0.15 mol m^?2 s^?1) above which estimates of F_A and F_R became larger in magnitude relative to GEE and TER. The value of D decreased with increasing values of g_m and g_cmax, but was still positive across all values of g_m and g_cmax. We conclude that the characterization of canopy‐scale mesophyll and stomatal conductances are important for further progress with the isotope partitioning method, and to confirm our anomalous isotopic disequilibrium findings.     

稻鸭共作系统的稻田氮素渗漏和径流特征

通过田间试验,分析了稻鸭共作（MRD）、常规栽培(MR)和淹灌单作（CK）3种稻作方式的稻田氮素渗漏和径流特征.结果表明：与MR处理相比,稻鸭共作系统稻田渗漏液氮素浓度尤其是NO₃^--N浓度显著减少,与CK处理相比,稻鸭共作稻田渗漏液氮素浓度有增加趋势;MRD处理在施肥7～9 d后,田面水中的NH₄⁺-N和NO₃^--N浓度高于MR处理,由于MRD处理不用搁田且田埂较高,田间排水量显著减少,氮素径流损失反而显著少于MR处理;与MR处理相比,MRD处理增加了鸭粪的氮素投入,减少了氮素渗漏和径流损失,减少了化学氮肥用量,增加了水稻地上部吸氮量;MRD系统的氮素输入总量和氮素输出总量均减少,且减少量基本相等.     

Photosynthesis as related to xylem water potential and carbon dioxide concentration in Sitka spruce

Current-year shoots of Sitka spruce ( Picea sitchensis (Bong.) Carr.) were removed from the forest canopy. After steady-state rates of net photosynthesis were obtained in a leaf chamber, the shoots were excised in air and removed at different times to establish a relationship between net photosynthesis and xylem water potential. The experiment was repeated at five ambient carbon dioxide concentrations.
Net photosynthesis remained constant over a wide range of xylem water potential and increased linearly with ambient carbon dioxide concentration between 20 and 300 cm³ m⁻³. At low water potential net photosynthesis declined at each ambient carbon dioxide concentration and there was little difference in the potential (±0.05 MPa) at which zero photosynthesis was observed.
There was a small increase in the CO₂ compensation concentration at low xylem water potentials, but calculated mesophyll conductance still declined at low water potential after correction for this change in compensation concentration. Mesophyll conductance reached zero within the same range of water potential as net photosynthesis. The results suggested that the non-stomatal contribution to the decline of photosynthesis was approximately 30% until almost complete stomatal closure occurred.     

To concentrate or ventilate? Carbon acquisition,isotope discrimination and physiological ecology of early land plant life forms

A comparative study has been made of the photosynthetic physiological ecology and carbon isotope discrimination characteristics for modern-day bryophytes and closely related algal groups. Firstly, the extent of bryophyte distribution and diversification as compared with more advanced land plant groups is considered. Secondly, measurements of instantaneous carbon isotope discrimination (Delta), photosynthetic CO(2) assimilation and electron transport rates were compared during the drying cycles. The extent of surface diffusion limitation (when wetted), internal conductance and water use efficiency (WUE) at optimal tissue water content (TWC) were derived for liverworts and a hornwort from contrasting habitats and with differing degrees of thallus ventilation (as intra-thalline cavities and internal airspaces). We also explore how the operation of a biophysical carbon-concentrating mechanism (CCM) tempers isotope discrimination characteristics in two other hornworts, as well as the green algae Coleochaete orbicularis and Chlamydomonas reinhardtii. The magnitude of Delta was compared for each life form over a drying curve and used to derive the surface liquid-phase conductance (when wetted) and internal conductance (at optimal TWC). The magnitude of external and internal conductances, and WUE, was higher for ventilated, compared with non-ventilated, liverworts and hornworts, but the values were similar within each group, suggesting that both factors have been optimized for each life form. For the hornworts, leakiness of the CCM was highest for Megaceros vincentianus and C. orbicularis (approx. 30%) and, at 5%, lowest in C. reinhardtii grown under ambient CO2 concentrations. Finally, evidence for the operation of a CCM in algae and hornworts is considered in terms of the probable role of the chloroplast pyrenoid, as the origins, structure and function of this enigmatic organelle are explored during the evolution of land plants.     

耦合叶肉导度的陆面过程模型最大叶肉导度参数的敏感性分析

陆面过程模型添加叶肉导度能有效改善模型模拟的CO₂施肥效应精度,但叶肉导度模拟受最大叶肉导度参数取值的影响,优化模型中最大叶肉导度参数是改进陆面过程模型叶肉导度和CO₂施肥效应模拟的重要途径。以EALCO（Ecological Assimilation of Land and Climate Observations）模型为例添加叶肉导度,通过人为改变最大叶肉导度值的取值,分析模型输出结果对最大叶肉导度的响应,揭示最大叶肉导度参数在模型中的敏感性,并与已有研究结果或观测数据比较,探讨耦合叶肉导度的陆面过程模型最大叶肉导度参数优化的途径。模拟试验以美国哈佛森林典型温带落叶阔叶林生态监测站（US-Ha1 site, Harvard Forest Environmental Monitoring site）数据为驱动。结果显示：（1）随最大叶肉导度增加,总初级生产力（GPP, Gross Primary Production）模拟精度增加,但最大叶肉导度取值大于1.0 mol m^-2 s^-1后模拟精度改善有限,最大叶肉导度小于1.0 mol m^-2 s^-1时GPP模拟精度对最大叶肉导度变化响应敏感;（2）证实了叶肉导度与气孔导度之间存在明显线性关系,最大叶肉导度取值的变化能明显影响这种线性关系的斜率。当最大叶肉导度取值从0.5 mol m^-2 s^-1增加到1.2 mol m^-2 s^-1时,气孔导度与叶肉导度的比值从0.75左右降至0.36,这个结果表明,通过明确某一植被功能型叶肉导度与气孔导度比值,可以间接确定模型最大叶肉导度的合理取值范围;（3）证实了陆面过程模型添加叶肉导度能改进CO₂施肥效应模拟精度,最大叶肉导度值能影响施肥效应模拟结果,当最大叶肉导度高于0.57 mol m^-2 s^-1后,随最大叶肉导度增加,模拟GPP随大气CO₂浓度增加的增长率呈下降趋势;（4）在月尺度上叶肉导度模拟对最大叶肉导度值的敏感性随不同生长季而不同,在生长盛期的7、8月份最大叶肉导度对叶肉导度模拟结果影响最大,其次是5、6、9月份等生长次盛期,其他月份的影响较小。