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。     

采用稳定碳同位素法分析白羊草在不同干旱胁迫下的水分利用效率

本研究以黄土高原乡土草种白羊草(Bothriochloa ischaemum(L.)Keng.)为研究对象,采用盆栽控制实验,比较白羊草在3个水分处理(CK80%FC、MS60%FC和SS 40%FC)下的生物量积累和分配模式、瞬时水分利用效率(WUE)、不同部位(新叶、老叶、茎、细根、粗根)的稳定碳同位素组成(δ~(13)C)和碳同位素分辨率(Δ~(13)C)及其相互关系,以及干旱胁迫下影响水分利用效率的主导环境因子。结果表明:1)重度干旱胁迫显著降低植物整体生物量,显著增加根冠比和细根生物量比例;2)随着干旱胁迫加剧,白羊草各器官的δ~(13)C均呈上升趋势,Δ~(13)C呈减小趋势,SS处理不同器官δ~(13)C和Δ~(13)C没有显著差异,CK和MS处理的各器官δ~(13)C均值表现分别为细根粗根老叶新叶茎、细根新叶老叶粗根茎,CK和MS处理Δ~(13)C的值总体呈根叶茎。3)新叶的δ~(13)CNL和Δ~(13)CNL与WUE的相关系数均最大,说明利用稳定碳同位素方法测定白羊草水分利用效率具有可行性。4)不同水分处理的WUE的主导影响因子不同,CK、MS、SS水分处理WUE分别受到叶面温度、大气水汽压亏缺和空气温度的影响最大。为采用稳定碳同位素方法指示白羊草水分利用效率可行性及阐明植物的胁迫响应机制提供理论依据。     

荒漠植物叶片或同化枝δ13C值及水分利用效率研究

温带荒漠植物长期生长在夏季炎热高温、冬季寒冷低温、长年干旱缺水的极端环境中,以其特有的形态学特征和生理功能来减少水分损失.为了探讨荒漠植物水分利用效率(WUE),为荒漠生态系统保育和退化植被恢复重建提供理论依据,运用稳定碳同位素技术和光合仪测定对比的方法,对河西走廊中段临泽绿洲北部荒漠中5种植物进行了全年生长期研究.结果表明荒漠植物月水分利用效率与年生长期平均水分利用效率的相关性在8月份最高,其方程式为WUEgpa= -1.8 + 1.98 WUEAug(P = 0.011,r =0.96);月稳定碳同位素比率(δ13C 或13 C/12C) (‰)与生长期平均WUE的相关性在8月和9月最大,可靠性最高,其方程式为WUE gpa= 4.7 + 0.0813 C/12Cmon(P=0.057,r=0.87).研究得出用稳定碳同位素比率指示温带荒漠植物的短期水分利用效率,随着叶片或同化枝成熟,越往生长后期,正相关性越高,直至霜降;用稳定碳同位素比率指示植物的长期水分利用效率,以8月下旬至9月下旬采样最好.梭梭和沙拐枣的水分利用效率显著高于其它荒漠植物,5种荒漠植物长期水分利用效率的排列顺序为梭梭＞沙拐枣＞柠条＞花棒≈泡泡刺.     

稳定碳同位素测定水分利用效率——以决明子为例

通过称重法和稳定碳同位素方法对盆栽决明子生长不同时期不同部位(根、茎、叶)的碳同位素组成(δ13C)、碳同位素分辨率(Δ)和水分利用效率(WUE)进行了研究,并分析了它们之间的相关关系,结果表明:决明子不同部位δ13C值表现为根茎叶,Δ为根茎叶,不同时期方差分析显示只有根部差异显著,茎叶的δ13C值、碳同位素分辨率分别与水分利用效率呈显著的正相关和负相关关系。称重法得出的WUE和不同部位(根、茎、叶)碳同位素测定的水分利用效率(WUER、WUES、WUEL)有显著正线性相关,其中与WUEL相关系数最大为0.86,与WUES、WUER的相关系数分别为0.80和0.82。说明利用稳定碳同位素方法测定决明子水分利用效率具有可行性,尤其是利用叶片测定的水分利用效率得到更为可靠的结果。     

环境条件对植物稳定碳同位素组成的影响

植物稳定碳同位素技术是近年兴起的一项快速、可靠的技术。利用稳定碳同位素技术可以揭示碳同化的过程的许多方面的信息。1 3C和1 2 C同位素效应 ,使它们在进行碳循环时发生稳定碳同位素的分馏。植物光合作用过程中CO2 经气孔扩散分差和RUBPCase及PEPCase羧化分馏是造成植物稳定碳同位素比率 (R =1 3C/ 1 2 C)不同于源CO2 中碳同位素比率的主要原因。遗传因素和环境因子同时决定植物碳同位素组成。植物稳定碳同位素技术同时还是古气候重建和预测未来环境变化的理论基础。本文综述了光照、温度、水分、二氧化碳、矿质营养、盐分和大气污染物等环境因素对植物稳定碳同位素组成影响方面的研究进展。     

环境条件对植物稳定碳同位素组成的影响

植物稳定碳同位素技术是近年兴起的一项快速、可靠的技术。利用稳定碳同位素技术可以揭示碳同化的过程的许多方面的信息。13C和12C同位素效应,使它们在进行碳循环时发生稳定碳同位素的分馏。植物光合作用过程中CO2经气孔扩散分差和RUBPCase及PEPCase羧化分馏是造成植物稳定碳同位素比率(R=13C/12C)不同于源CO2中碳同位素比率的主要原因。遗传因素和环境因子同时决定植物碳同位素组成。植物稳定碳同位素技术同时还是古气候重建和预测未来环境变化的理论基础。本文综述了光照、温度、水分、二氧化碳、矿质营养、盐分和大气污染物等环境因素对植物稳定碳同位素组成影响方面的研究进展。     

水分和氮磷水平对小麦碳同位素分辨率和水分利用效率的影响

 依据盆栽试验数据,利用植物稳定性碳同位素分辨率的理论模型,研究了水分和氮磷营养对小麦叶片碳同位素分辨率(Carbon-isotope discrimination △)的影响。结果表明：水分差异引起碳同位素分辨率较大变异,碳同位素分辨率随土壤相对含水量(Soil relative water content)的提高而提高,在土壤相对含水量为60％～70％条件下碳同位素分辨率最高。缺水时磷水平提高,碳同位素分辨率提高。水分利用效率(Water use efficiency)与碳同位素分辨率关系受土壤水分和养分水平的影响。缺水条件下水分利用效率与碳同位素分辨率之间为负相关,充分供水下为正相关;在低氮水平下的关系不明显,施氮150kg·hm-2时相关性显著。     

毛乌素沙地三种灌木群落的水分利用过程

沙地柏、黑沙蒿和沙柳是毛乌素沙地的3种优势灌木群落。利用稳定同位素技术研究了3种灌木及伴生植物杨柴主要利用的水分来源,结合叶片稳定碳同位素值与土壤水分监测,从而确定灌木群落如何利用水分。结果表明:7月和9月3种群落内浅层土壤水的稳定氧同位素值接近雨水。沙地柏5月主要利用25 cm浅层土壤水,而7月和9月主要利用10—25 cm浅层和100—200 cm深层土壤水。黑沙蒿和伴生的杨柴5月主要利用10 cm浅层土壤水,7月同时利用10 cm浅层土壤水和150 cm深层土壤水,9月则利用10—150 cm土壤水。沙柳5月主要利用10—25 cm浅层土壤水,伴生的杨柴主要利用50—200 cm土壤水;7月它们同时利用10—25 cm浅层土壤水和100—200 cm深层土壤水;9月都主要利用25—200 cm土壤水。4种植物的叶片稳定碳同位素值存在季节动态和种间差异。常绿灌木沙地柏的叶片碳同位素值比较稳定,而且高于其他3种落叶灌木和半灌木。5月浅层土壤含水量较低时3种落叶植物的叶片碳同位素值较高。因此,降雨补充的浅层土壤水是3种灌木群落利用的主要水分来源。3种灌木及其伴生植物根据不同深度土壤水的可利用性,在不同季节利用不同深度的土壤水。杨柴与黑沙蒿或沙柳均存在水分竞争。沙地柏的叶片稳定碳同位素值较高,干旱时具有竞争优势。干旱时落叶灌木和半灌木能够提高叶片稳定碳同位素值来适应环境。     

稳定性碳同位素在植物生理生态研究中的应用

稳定性碳同位素~(13)C/~(12)C比率是研究植物光合作用及有关的物质代谢、水分关系和生态系统中种间关系、进化及对环境的响应等的有效指标。δ~(13)C值与细胞间CO_2浓度及水分利用效率之间有一定的数量关系。光合作用过程中对碳同位素的辨别力主要是不同的CO_2扩散阻抗和羧化反应速率引起的结果。     

基于稳定同位素的干旱半干旱区SPAC水分运移过程研究进展

土壤-植物-大气连续体(SPAC)是生态水文学的重点研究对象,其水分运移过程对于干旱半干旱区生态植被建设和水资源综合管理具有重要意义。氢氧稳定同位素较高的灵敏性和准确度有助于揭示这一过程。介绍了氢氧稳定同位素在土壤-大气界面、土壤-地下水界面、土壤-植物界面和植物-大气界面水分补给传输过程中的应用,包括土壤水分来源和蒸发;水分补给入渗机制和滞留时间;植物水分来源和水力再分配;蒸散发分割和叶片吸水的相关研究,同时明确了氢氧稳定同位素技术在应用过程中存在的一些不确定性以及未来亟需加强的方面,以期为利用稳定同位素技术对生态水文过程的研究提供参考依据。     

Stable carbon isotopes as indicators of increased water use efficiency and productivity in white spruce (Picea glauca (Moench) Voss) seedlings

The relationship among water use efficiency (WUE), productivity and carbon isotopic composition (δ¹³C) in white spruce (Picea glauca (Moench) Voss) seedlings was investigated. Sixteen hundred seedlings representing 10 controlled crosses were planted in the field in individual buried sand-filled cylinders. The soil water content in the cylinders was measured using time domain reflectometry over two growing seasons and seedling water use determined by water balance. Two watering treatments were imposed: irrigation and dry land. There was significant (1.6–2.0%c) genetic variation in needle δ¹³C. Ranking of crosses in terms of δ¹³C was generally maintained over watering treatments and there was not a significant genetic versus environmental interaction. There was a positive correlation between δ¹³C and both intrinsic and long-term WUE (more positive δ¹³C with increased WUE) and between δ¹³C and productivity, suggesting a correlation due to variation in photosynthetic capacity. Root to shoot ratios did not increase in water-stressed plants, indicating that responses to drought were primarily at the level of gas exchange, rather than through morphological changes. Our results indicate that it should be possible to use δ¹³C as a surrogate for WUE and to select white spruce genotypes for high WUE without compromising yield.     

Carbon isotope fractionation in tree ring early and late wood in relation to intra-growing season water balance

We determined the stable carbon isotope composition (δ^1.3C) of cellulose extracted from early and late wood in Douglas fir [Pseudotsuga menziexii (Mirb.) Franco] tree rings. Data were obtained for the period 1962 to 1981, at the start of which the trees were 20 years old. A water balance model was used to calculate daily stand transpiration and water deficit. The model incorporates site factors (soil water availability, slope and aspect) and environmental variables (solar radiation, air temperature and rainfall). There was far greater variability in late wood than in early wood δ^1.3C. In wet years, late wood δ^1.3C was significantly lighter (by as much as 2δ) than early wood δ^1.3C but in dry years this difference was reversed. Differences between spring and summer cumulative transpiration accounted for almost 60δ of the variability in differences between early and late wood δ^1.3C. We found excellent correspondence between summer cumulative transpiration and late wood δ^1.3C, with estimates of transpiration accounting for up to 93% of the variability in δ^1.3C. Correlations between early wood δ^1.3C and spring transpiration were generally poor (r2<0.4), but we were able to identify those exceptional years in which there had been a very dry spring. Our results indicate that, while tree ring δ^1.3C correlates reasonably well with basal area increment, it is a far better indicator of inter- and intra-annual variability in water availability than radial growth.     

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.     

Does the 13C of foliage‐respired CO2 and biochemical pools reflect the 13C of recently assimilated carbon?

Isotopic labelling experiments were conducted to assess relationships among ¹³C of recently assimilated carbon ( δC _A), foliage respiration ( δC _F), soluble carbohydrate ( δC _SC), leaf waxes ( δC _LW) and bulk organic matter ( δC _OM). Slash pine, sweetgum and maize were grown under ¹³C depleted CO₂ to label biomass and then placed under ambient conditions to monitor the loss of label. In pine and sweetgum, δC _F of labelled plants (∼−44 and −35‰, respectively) rapidly approached control values but remained depleted by ∼4–6‰ after 3–4 months. For these tree species, no or minimal label was lost from δC _SC, δC _LW and δC _OM during the observation periods. δC _F and δC _SC of labelled maize plants rapidly changed and were indistinguishable from controls after 1 month, while δC _LW and δC _OM more slowly approached control values and remained depleted by 2–6‰. Changes in δC _F in slash pine and sweetgum fit a two-pool exponential model, with the fast turnover metabolic pool (∼3–4 d half-life) constituting only 1–2% of the total. In maize, change in δC _F fits a single pool model with a half-life of 6.4 d. The ¹³C of foliage respiration and biochemical pools reflect temporally integrated values of δC _A, with change in isotopic composition dampened by the size of metabolic carbon reserves and turnover rates.     

Dietary analysis of the herbivorous hemiramphid Hyporhamphus regularis ardelio: an isotopic approach

The stable isotope values for a range of size classes of Hyporhamphus regularis ardelio from Moreton Bay, south‐east Australia were determined. There was a positive linear relationship between δ¹³C and standard length (L_S)(δ¹³C = 0·034 L_S ? 16·23; r² = 0·78). δ¹³C ranged from ?8·48 to ?17·29‰ with the smallest size class (50 mm L_S) being on average 1·04‰ enriched with respect to that of zooplankton (Temora turbinata) and 7·97‰ depleted compared to Zostera capricorni. δ¹³C was positively correlated with L_S(P < 0·01)(more enriched with increasing L_S) with those fish of the largest size class (225 mm L_S) being 9·86 and 0·84‰ enriched than T. turbinata and Z. capricorni, respectively. There was no detectable trend in δ¹⁵N values with L_S(P > 0·01) with δ¹⁵N, ranging from 9·18 to 11·00‰. Fish of all size classes were on average 2·32 and 7·63‰ more enriched than zooplankton and seagrass, respectively. Carbon isotope data indicate that H. r. ardelio commence life as carnivores and change to a diet in which seagrass is the primary carbon source. The dependence on animal matter, however, is always present. Due to the low percentage of nitrogen in Z. capricorni(2·5%) compared to zooplankton (9·1%) it appears that nitrogen from zooplankton is necessary throughout their life history with the carbon requirements for these fish coming chiefly from Z. capricorni.     

The effects of nitrogen stress on the stable carbon isotope composition, productivity and water use efficiency of white spruce (Picea glauca (Moench) Voss) seedlings

A, assimilation rate
a, fractionation against ¹³C for CO₂ diffusion through air
b, net fractionation against ¹³C during CO₂ fixation
C_a, ambient CO₂ concentration
C_c, CO₂ concentration at the chloroplast
C_i, intercellular CO₂ concentration
D, vapour pressure deficit
E_n, needle transpiration rate
E_p, whole plant water use
g_w, leaf internal transfer conductance to CO₂
g_s, stomatal conductance to water vapour
L, projected leaf area
NUE, nitrogen use efficiency
PEP, phosphoenolpyruvate
Rubisco, ribulose-1,5-biphosphate carboxylase
TDR, time domain reflectometry
WUE, water use efficiency
Δ, carbon isotope discrimination
δ¹³C, carbon isotope abundance parameter
δ¹³C_a, carbon isotopic composition of atmospheric CO₂
θ, volumetric soil water content

The effect of nitrogen stress on needle δ¹³C, water-use efficiency (WUE) and biomass production in irrigated and dry land white spruce (Picea glauca (Moench) Voss) seedlings was investigated. Sixteen hundred seedlings, representing 10 controlled crosses, were planted in the field in individual buried sand-filled cylinders. Two nitrogen treatments were imposed, nitrogen stressed and fertilized. The ranking of δ¹³C of the crosses was maintained across all combinations of water and nitrogen treatments and there was not a significant genetic versus environmental interaction. The positive relationships between needle δ¹³C, WUE and dry matter production demonstrate that it should be possible to use δ¹³C as a surrogate for WUE, and to select for increased WUE without compromising yield, even in nitrogen deficient environments. Nitrogen stressed seedlings had the lowest needle δ¹³C in both irrigated and dry land conditions. There was a positive correlation between needle nitrogen content and δ¹³C that was likely associated with increased photosynthetic capacity. There was some indication that decreased nitrogen supply led to increased stomatal conductance and hence lower WUE. There was a negative correlation between intrinsic water use efficiency and photosynthetic nitrogen use efficiency (NUE). This suggests that white spruce seedlings have the ability to maximize NUE when water becomes limited. There was significant genetic variation in NUE that was maintained across treatments. Our results suggest that in white spruce, there is no detectable effect of anaplerotic carbon fixation and that it is more appropriate to use a value of 29‰ (‘Rubisco only’) for the net discrimination against ¹³C during CO₂ fixation. This leads to excellent correspondence between values of C_i/C_a derived from gas exchange measurements or from δ¹³C.     

多年放牧对不同类型草原植被及土壤碳同位素的影响

放牧是草原牧区常见的人类活动,多年放牧对草原植被及土壤的碳过程产生较大的影响.本研究采集不同类型草原多年放牧前后植被及土壤样品,对室内碳同位素进行分析,研究了不同草原生态系统Δ¹³C(碳同位素分馏值)差异及其影响因素.结果表明: 放牧强度对植被Δ¹³C值的影响显著,0～5 cm表层土壤Δ¹³C值在放牧前后变化显著,而对深层土壤(>5 cm)影响不显著.多年放牧后大部分植被Δ¹³C值显著升高,高海拔地区升高的幅度较大.可见,放牧行为对不同草地生态系统类型、不同土壤深度以及不同海拔生态系统碳过程产生的影响差异显著.针对不同类型的草原,放牧应采取多样化的管理方式.