植物叶片水稳定同位素研究进展

植物叶片水稳定同位素变化可以直接沟通植物叶片内部与外界的物质和能量联系,并能够反映植物生长周围的气候与生态信息.另外,植物叶片水作为参与水循环的一个重要环节,了解叶片水稳定同位素组成有助于揭示其在局地水体稳定同位素循环中的分配与贡献.概述了国内外叶片水稳定同位素研究现况;介绍了叶片水稳定氢、氧同位素在植物体中的分馏过程及形式(热力学平衡分馏、动力学分馏以及生化分馏)以及影响叶片水稳定同位素组成的气象和生态因子;阐述了叶片水稳定同位素修正的Craig-Gordon稳态模型、string-of-lakes模型、Péclet效应的稳态模型、非稳态效应的模型、Péclet效应的非稳态模型以及二维模型的构建与完善过程;最后讨论了植物叶片水稳定同位素研究存在的问题,并从叶片水稳定同位素与气象、生态因子的关系,叶片水蒸腾线的斜率和截距及过量氘的意义,模型适用性的验证以及叶片水稳定同位素在水文循环的应用等方面展望了研究方向.     

Modelling the isotope enrichment of leaf water

Farquhar and Gan [10] have proposed a model for the spatial variation in the isotopic enrichment of H₂¹⁸O across a leaf, which is specifically formulated for monocotyledoneous leaves. The model is based on the interaction between mass fluxes longitudinally within the xylem, and fluxes laterally through veinlets into the lamina mesophyll, where moisture leaves the leaf through transpiration. The lighter, more abundant, molecule H₂¹⁶O escapes preferentially with the evaporating water, resulting in the enrichment of H₂¹⁸O at these sites. Enriched water diffuses throughout the leaf, and it is this spatial distribution of enriched water which the model seeks to capture. In this paper we present a general formulation of the model in terms of mass flux, extending it to include variable transpiration rates across the leaf surface, as well as a tapering xylem. Solutions are developed for the general case and, since the solutions present in the form of Kummer functions, properties are established as well as methods for estimating the solutions under certain conditions relevant to the biology. The model output is compared with Gans data ([14, 15]) collected from maize plants.     

Effects of environmental parameters, leaf physiological properties and leaf water relations on leaf water delta18O enrichment in different Eucalyptus species

Stable oxygen isotope ratios (delta18O) have become a valuable tool in the plant and ecosystem sciences. The interpretation of delta18O values in plant material is, however, still complicated owing to the complex interactions among factors that influence leaf water enrichment. This study investigated the interplay among environmental parameters, leaf physiological properties and leaf water relations as drivers of the isotopic enrichment of leaf water across 17 Eucalyptus species growing in a common garden. We observed large differences in maximum daily leaf water delta18O across the 17 species. By fitting different leaf water models to these empirical data, we determined that differences in leaf water delta18O across species are largely explained by variation in the Péclet effect across species. Our analyses also revealed that species-specific differences in transpiration do not explain the observed differences in delta18O while the unconstrained fitting parameter 'effective path length' (L) was highly correlated with delta18O. None of the leaf morphological or leaf water related parameters we quantified in this study correlated with the L values we determined even though L was typically interpreted as a leaf morphological/anatomical property. A sensitivity analysis supported the importance of L for explaining the variability in leaf water delta18O across different species. Our investigation highlighted the importance of future studies to quantify the leaf properties that influence L. Obtaining such information will significantly improve our understanding of what ultimately determines the delta18O values of leaf water across different plant species.     

Variability in mesophyll conductance between barley genotypes,and effects on transpiration efficiency and carbon isotope discrimination

Leaf internal, or mesophyll, conductance to CO₂ (g_m ) is a significant and variable limitation of photosynthesis that also affects leaf transpiration efficiency (TE). Genotypic variation in g_m and the effect of g_m on TE were assessed in six barley genotypes (four Hordeum vulgare and two H. bulbosum). Significant variation in g_m was found between genotypes, and was correlated with photosynthetic rate. The genotype with the highest g_m also had the highest TE and the lowest carbon isotope discrimination as recorded in leaf tissue (Δ_p). These results suggest g_m has unexplored potential to provide TE improvement within crop breeding programmes.     

A dynamic leaf gas‐exchange strategy is conserved in woody plants under changing ambient CO2: evidence from carbon isotope discrimination in paleo and CO2 enrichment studies

Rising atmospheric [CO₂], c_a, is expected to affect stomatal regulation of leaf gas‐exchange of woody plants, thus influencing energy fluxes as well as carbon (C), water, and nutrient cycling of forests. Researchers have proposed various strategies for stomatal regulation of leaf gas‐exchange that include maintaining a constant leaf internal [CO₂], c_i, a constant drawdown in CO₂ (c_a ? c_i), and a constant c_i/c_a. These strategies can result in drastically different consequences for leaf gas‐exchange. The accuracy of Earth systems models depends in part on assumptions about generalizable patterns in leaf gas‐exchange responses to varying c_a. The concept of optimal stomatal behavior, exemplified by woody plants shifting along a continuum of these strategies, provides a unifying framework for understanding leaf gas‐exchange responses to c_a. To assess leaf gas‐exchange regulation strategies, we analyzed patterns in c_i inferred from studies reporting C stable isotope ratios (δ¹³C) or photosynthetic discrimination (?) in woody angiosperms and gymnosperms that grew across a range of c_a spanning at least 100 ppm. Our results suggest that much of the c_a‐induced changes in c_i/c_a occurred across c_a spanning 200 to 400 ppm. These patterns imply that c_a ? c_i will eventually approach a constant level at high c_a because assimilation rates will reach a maximum and stomatal conductance of each species should be constrained to some minimum level. These analyses are not consistent with canalization toward any single strategy, particularly maintaining a constant c_i. Rather, the results are consistent with the existence of a broadly conserved pattern of stomatal optimization in woody angiosperms and gymnosperms. This results in trees being profligate water users at low c_a, when additional water loss is small for each unit of C gain, and increasingly water‐conservative at high c_a, when photosystems are saturated and water loss is large for each unit C gain.     

Associations between carbon isotope ratios of ecosystem respiration, water availability and canopy conductance

We tested the hypothesis that the stable carbon isotope signature of ecosystem respiration (δ¹³C_R) was regulated by canopy conductance (G_c) using weekly Keeling plots (n=51) from a semiarid old‐growth ponderosa pine (Pinus ponderosa) forest in Oregon, USA. For a comparison of forests in two contrasting climates we also evaluated trends in δ¹³C_R from a wet 20‐year‐old Douglas‐fir (Pseudotsuga menziesii) plantation located near the Pacific Ocean. Intraannual variability in δ¹³C_R was greater than 8.0‰ at both sites, was highest during autumn, winter, and spring when rainfall was abundant, and lowest during summer drought. The δ¹³C_R of the dry pine forest was consistently more positive than the wetter Douglas‐fir forest (mean annual δ¹³C_R: ?25.41‰ vs. ?26.23‰, respectively, P=0.07). At the Douglas‐fir forest, δ¹³C_R–climate relationships were consistent with predictions based on stomatal regulation of carbon isotope discrimination (Δ). Soil water content (SWC) and vapor pressure deficit (vpd) were the most important factors governing δ¹³C_R in this forest throughout the year. In contrast, δ¹³C_R at the pine forest was relatively insensitive to SWC or vpd, and exhibited a smaller drought‐related enrichment (～2‰) than the enrichment observed during drought at the Douglas‐fir forest (～5‰). Groundwater access at the pine forest may buffer canopy–gas exchange from drought. Despite this potential buffering, δ¹³C_R at the pine forest was significantly but weakly related to canopy conductance (G_c), suggesting that δ¹³C_R remains coupled to canopy–gas exchange despite groundwater access. During drought, δ¹³C_R was strongly correlated with soil temperature at both forests. The hypothesis that canopy‐level physiology is a critical regulator of δ¹³C_R was supported; however, belowground respiration may become more important during rain‐free periods.     

Daily variations in water relations of apricot trees under different irrigation regimes

Mature apricot (Prunus armeniaca L. cv. Búlida) trees, growing under field conditions, were submitted to two drip irrigation treatments: a control (T1), irrigated to 100 % of seasonal crop evapotranspiration (ETc), and a continuous deficit (T2), irrigated to 50 % of the control throughout the year. The behaviour of leaf water potential and its components, leaf conductance and net photosynthesis were studied at three different times during the growing season, when they revealed a diurnal and seasonal pattern in response to water stress, evaporative demand of the atmosphere and leaf age. The deficit-irrigated trees showed, among other effects, a pronounced decrease in leaf water potential (ψ_w), decreased in leaf conductance (g_s) and no osmotic adjustment. For this reason, g_l and ψ_w can be considered good indicators of mature apricot tree water status and can therefore be used for irrigation scheduling.     

Differential coordination of stomatal conductance,mesophyll conductance,and leaf hydraulic conductance in response to changing light across species

Stomatal conductance (g_s) and mesophyll conductance (g_m) represent major constraints to photosynthetic rate (A), and these traits are expected to coordinate with leaf hydraulic conductance (K_leaf) across species, under both steady‐state and dynamic conditions. However, empirical information about their coordination is scarce. In this study, K_leaf, gas exchange, stomatal kinetics, and leaf anatomy in 10 species including ferns, gymnosperms, and angiosperms were investigated to elucidate the correlation of H₂O and CO₂ diffusion inside leaves under varying light conditions. Gas exchange, K_leaf, and anatomical traits varied widely across species. Under light‐saturated conditions, the A, g_s, g_m, and K_leaf were strongly correlated across species. However, the response patterns of A, g_s, g_m, and K_leaf to varying light intensities were highly species dependent. Moreover, stomatal opening upon light exposure of dark‐adapted leaves in the studied ferns and gymnosperms was generally faster than in the angiosperms; however, stomatal closing in light‐adapted leaves after darkening was faster in angiosperms. The present results show that there is a large variability in the coordination of leaf hydraulic and gas exchange parameters across terrestrial plant species, as well as in their responses to changing light.     

Stomatal response to leaf water potential in almond trees under drip irrigated and non irrigated conditions

Almond plants (Amygdalus communis L. cv. Garrigues) were grown in the field under drip irrigated and non irrigated conditions. Leaf water potential () and leaf conductance (g₁) were determined at three different times of the growing season (spring, summer and autumn). The relationships between and g₁ in both treatments showed a continuous decrease of g₁ as decreased in spring and summer. Data from the autumn presented a threshold value of (approx. –2.7 MPa in dry treatment, and approx. –1.4 MPa in wet treatment) below which leaf conductance remained constant.     

Photosynthetic CO2 uptake byZea mays leaves as influenced by unilateral irradiation of adaxial and abaxial leaf surfaces

A study was made on the effect of increasing photon fluence rate (I) at a unilateral irradiation of adaxial (normal leaf position) and abaxial (inverse leaf position) blade surface of maize leaves of various insertion levels on net photosynthetic CO₂ uptake (P _n ) by the leaves, as well as the contribution of individual surfaces toP _n of the leaves, and the significance of, or relationship between the stomatal (g _s ) and intracellular (gm) conductances at the CO₂ transport.P _n of leaves of various age according to their insertion level was unaffected by the direction of incident irradiation. Upon irradiation of the leaves in normal and inverse position the contribution of the adaxial and abaxial surfaces toP _n ,g _s and g_m was different. On irradiating the leaves in normal position, the contribution of the irradiated adaxial surface to the characteristics mentioned made on the average 55% of total values, the contribution of the abaxial surface irradiated in inverse position made on the average 70% inP _n andg _m , and 80% ing _s . At lowerI’s g _m was higher thang _s both in irradiated and non-irradiated surfaces. The ratio ofg _s to g_m gradually got square with increasingI. In the irradiated adaxial surface the equilibrium (g _s /g _m = 1.0) took place at the highestI’s, in the irradiated abaxial surface between 500 to 1000 μmol m⁻² s⁻¹. The significance of the ratiog _m in the CO₂ transport through the individual surfaces is discussed.     

Strategies of leaf expansion in Ficus carica under semiarid conditions

Leaf area expansion, thickness and inclination, gas exchange parameters and relative chlorophyll content were analysed in field‐grown fig (Ficus carica L.) leaves over time, from emergence until after full leaf expansion (FLE). Ficus carica leaves showed a subtle change in shape during the early stages of development, and FLE was reached within ca. 30 days after emergence. Changes in leaf thickness and inclination after FLE demonstrated good adaptation to environmental conditions during summer in areas with a Mediterranean climate. Changes in gas exchange parameters and relative chlorophyll content showed that F. carica is a delayed‐greening species, reaching maximum values 20 days after FLE. Correlation analysis of datasets collected during leaf expansion, confirmed dependence among structural and functional traits in F. carica. Pn was directly correlated with stomatal conductance (Gs), transpiration (E), leaf area (LA) and relative chlorophyll content up to FLE. The effect of pruning on leaf expansion, a cultural technique commonly applied in this fruit tree, was also evaluated. Although leaf development in pruned branches gave a significantly higher relative leaf area growth rate (RGR_l) and higher LA than non‐pruned branches, no significant differences were found in other morphological and physiological traits, indicating no pruning effect on leaf development. All studied morphological and physiological characteristics indicate that F. carica is well adapted to semiarid conditions. The delayed greening strategy of this species is discussed.     

Functional coordination between leaf gas exchange and vulnerability to xylem cavitation in temperate forest trees

We examined functional coordination among stem and root vulnerability to xylem cavitation, plant water transport characteristics and leaf traits in 14 co-occurring temperate tree species. Relationships were evaluated using both traditional cross-species correlations and phylogenetically independent contrast (PIC) correlations. For stems, the xylem tension at which 50% of hydraulic conductivity was lost (psi50) was positively associated (P < 0.001) with specific conductivity (K(S)) and with mean hydraulically weighted xylem conduit diameter (D(h-w)), but was only marginally (P = 0.06) associated with leaf specific conductivity (K(L)). The PIC correlation for each of these relationships, however, was not statistically significant. There was also no relationship between root psi50 and root K(S) in either cross-species or PIC analysis. Photosynthetic rate (A) and stomatal conductance (g(s)) were strongly and positively correlated with root psi50 in the cross-species analysis (P < 0.001), a relationship that was robust to phylogenetic correction (P < 0.01). A and g(s) were also positively correlated with stem psi50 in the cross-species analysis (P = 0.02 and 0.10, respectively). However, only A was associated with stem psi50 in the PIC analysis (P = 0.04). Although the relationship between vulnerability to cavitation and xylem conductivity traits within specific organs (i.e. stems and roots) was weak, the strong correlation between g(s) and root psi50 across species suggests that there is a trade-off between vulnerability to cavitation and water transport capacity at the whole-plant level. Our results were therefore consistent with the expectation of coordination between vulnerability to xylem cavitation and the regulation of stomatal conductance, and highlight the potential physiological and evolutionary significance of root hydraulic properties in controlling interspecific variation in leaf function.     

Photosynthesis, leaf conductance and water relations of in vitro cultured grapevine rootstock in relation to acclimatisation

Leaf net CO₂ uptake and leaf photosynthetic capacity were investigated in micropropagated 41B grapevine rootstock (Vitis vinifera‘Chasselas’×Vitis berlandieri, Mill. De Gr.) plants grown in the presence of four sucrose concentrations (6.25, 12.5, 25.0 or 37.5 g l^?1). Sucrose concentration in the medium during growth in vitro did not affect the leaf photosynthetic performance of plants neither before nor after transplantation. The maximum photosynthetic rate, measured as CO₂-dependent O₂ evolution, was 7.3 µmol m^?2 s^?1 before transplanting and 15.4 µmol m^?2 s^?1 one month after transplantation. The maximum quantum yield of O₂ evolution (on the basis of incident light) was about 0.07 for all sucrose treatments both before and after transplantation. Dry biomass before transplanting was highest in plants grown with 25.0 or 37.5 g l^?1 sucrose in the medium. One month after transplantation the highest dry biomass was also observed for the same treatments. Survival of plants was 100% for all treatments. Leaf conductance to water vapour was always higher in plants before than after transplantation. Both before and after transplanting it increased with increasing light intensity and decreased slightly with increasing CO₂ molar ratio in in vitro plants. Stomata of plants before transplantation were unresponsive to vapour pressure deficit. In vitro plants experience an acute water stress when they are maintained with the whole root system in water and exposed to ambient controlled conditions in a growth chamber. However, there was no wilting of the leaves when similar plants with roots cut off were left in the same conditions. Hydraulic conductivity was low at both root and shoot-root connection levels. It is likely that water supply could be limiting during transplantation because of the low root and root-stem connection conductivity. Water uptake by roots rather than water loss from the shoots would be of primary importance for the maintenance of water balance during acclimatisation.     

分根区施保水剂对玉米气孔导度和单叶WUE的影响

盆栽条件下,研究了陕单9号玉米(zea mays L．)在根区不施保水剂(对照)、分根区施保水剂和根区全施保水剂3种处理下,叶片气孔导度、CO2吸收和H2O蒸腾的变化。结果表明,在75％土壤饱和持水量下,各指标在3种处理之间没有明显差别;在50％土壤饱和持水量下,分根区施保水剂显著降低了叶片气孔导度,叶片CO2吸收量和H2O蒸腾量也同时降低,但H2O蒸腾量下降幅度更大;在两种水分条件下,分根区施保水剂均能提高玉米单叶水分利用效率(water use efficiency,WUE)。     

不同甘蔗品种叶片气孔对水分胁迫的响应

干旱是甘蔗面临最主要的环境胁迫之一,为了解不同甘蔗品种在干旱胁迫时的气孔响应,该研究以F172、GT21、YT93/159和 YL6四个抗旱性有显著差异的甘蔗品种为材料,采用桶栽,在伸长期进行四种不同程度的干旱胁迫(不浇水)处理：土壤持水量在①65％~70％为轻度干旱;②45％~50％为中度干旱;③25％~30％为重度干旱;④以土壤含水量为75％为对照(CK).检测不同品种不同处理甘蔗的叶片相对持水量变化,并利用扫描电镜技术观察甘蔗叶片下表皮气孔特性.结果表明：在干旱胁迫下,四个甘蔗品种叶片气孔导度急剧下降,重度干旱时耐旱性强的 F172和 GT21的气孔导度低于耐旱性弱的 YT93/159和 YL6的;复水后3 d,F172和 GT21的气孔导度上升至82．07和88．85 mmol·m-2·s-1,而 YT93/159和 YL6的仅有18．88和33．08 mmol·m-2·s-1.干旱还导致气孔下陷、闭合,气孔器的长、宽明显减小,且品种间气孔器长度变化差异显著;干旱胁迫下气孔密度增大,尤以耐旱性最强的 F172在重度干旱时达到显著差异.重度干旱时 F172与GT21的气孔闭合百分比是 YT93/159和 YL6近3~4倍.在水分胁迫下,叶片相对含水量降低,但 F172和GT21在重度干旱时仍可以保持相对较高的含水量,其它两个品种相对较低,尤以 YT93/159的最低.在复水后叶片含水量都有所恢复.这些研究结果表明不同甘蔗品种抗旱能力与叶片气孔特性和含水量密切相关.     

Non-steady state effects in diurnal 180 discrimination by Picea sitchensis branches in the field

We report diurnal variations in 18O discrimination (18 delta) during photosynthesis (18 delta A) and respiration (18 delta R) of Picea sitchensis branches measured in branch chambers in the field. These observations were compared with predicted 18 delta (18 delta pred) based on concurrent measurements of branch gas exchange to evaluate steady state and non-steady state (NSS) models of foliage water 18O enrichment for predicting the impact of this ecosystem on the Delta 18O of atmospheric CO2. The non-steady state approach substantially improved the agreement between 18 delta pred and observed 18 delta (18 delta obs) compared with the assumption of isotopic steady state (ISS) for the Delta 18O signature of foliage water. In addition, we found direct observational evidence for NSS effects: extremely high apparent 18 delta values at dusk, dawn and during nocturnal respiration. Our experiments also show the importance of bidirectional foliage gas exchange at night (isotopic equilibration in addition to the net flux). Taken together, neglecting these effects leads to an underestimation of daily net canopy isofluxes from this forest by up to 30%. We expect NSS effects to be most pronounced in species with high specific leaf water content such as conifers and when stomata are open at night or when there is high relative humidity, and we suggest modifications to ecosystem and global models of delta 18O of CO2.     

Photosynthetic gas exchange and the stable isotope composition of leaf water: comparison of a xylem-tapping mistletoe and its host

Photosynthetic gas exchange and the stable isotopic composition of foliage water were measured for a xylem tapping mistletoe, Phoradendron juniperinum, and its host tree, Juniperus osteosperma, growing in southern Utah. The observed isotopic composition of water extracted from foliage was compared to predictions of the Craig-Gordon model of isotopic enrichment at evaporative sites within leaves. Assimilation rates of juniper were higher and stomatal conductance was lower than the values observed for the mistletoe. This resulted in lower intercellular/ ambient CO₂ values in the juniper tree relative to its mistletoe parasite. For mistletoe, the observed foliage water hydrogen and oxygen isotopic enrichment was less than that predicted by the model. In juniper, foliage water hydrogen isotopic enrichment was also lower than that predicted by the evaporative enrichment model. In contrast, the oxygen isotopic enrichment in juniper foliage water was slightly greater than that predicted for the evaporative sites within leaves. Hydrogen isotopic enrichment in mistletoe foliage shows systematic variation with stem segment, being highest near the tips of the youngest stems and decreasing toward the base of the mistletoe, where isotopic composition is close to that of stem water in the host tree. In a correlated pattern, mid-day stomatal conductance declined abruptly in mistletoe foliage of increasing age.     

Impacts of tree height on leaf hydraulic architecture and stomatal control in Douglas-fir

This study investigated the mechanisms involved in the regulation of stomatal closure in Douglas-fir and evaluated the potential impact of compensatory adjustments in response to increasing tree height upon these mechanisms. In the laboratory, we measured leaf hydraulic conductance (K(leaf)) as leaf water potential (Psi(l)) declined for comparison with in situ diurnal patterns of stomatal conductance (g(s)) and Psi(l) in Douglas-fir across a height gradient, allowing us to infer linkages between diurnal changes in K(leaf) and g(s). A recently developed timed rehydration technique was used in conjunction with data from pressure-volume curves to develop hydraulic vulnerability curves for needles attached to small twigs. Laboratory-measured K(leaf) declined with increasing leaf water stress and was substantially reduced at Psi(l) values of -1.34, -1.45, -1.56 and -1.92 MPa for foliage sampled at mean heights of approximately 20, 35, 44 and 55 m, respectively. In situ g(s) measurements showed that stomatal closure was initiated at Psi(l) values of -1.21, -1.36, -1.74 and -1.86 MPa along the height gradient, which was highly correlated with Psi(l) values at loss of K(leaf). Cryogenic scanning electron microscopy (SEM) images showed that relative abundances of embolized tracheids in the central vein increased with increasing leaf water stress. Leaf embolism appeared to be coupled to changes in g(s) and might perform a vital function in stomatal regulation of plant water status and water transport in conifers. The observed trends in g(s) and K(leaf) in response to changes in Psi(l) along a height gradient suggest that the foliage at the tops of tall trees is capable of maintaining stomatal conductance at more negative Psi(l). This adaptation may allow taller trees to continue to photosynthesize during periods of greater water stress.     

A model explaining genotypic and ontogenetic variation of leaf photosynthetic rate in rice (Oryza sativa) based on leaf nitrogen content and stomatal conductance

BACKGROUNDS AND AIMS: Identification of physiological traits associated with leaf photosynthetic rate (Pn) is important for improving potential productivity of rice (Oryza sativa). The objectives of this study were to develop a model which can explain genotypic variation and ontogenetic change of Pn in rice under optimal conditions as a function of leaf nitrogen content per unit area (N) and stomatal conductance (g(s)), and to quantify the effects of interaction between N and g(s) on the variation of Pn. METHODS: Pn, N and g(s) were measured at different developmental stages for the topmost fully expanded leaves in ten rice genotypes with diverse backgrounds grown in pots (2002) and in the field (2001 and 2002). A model of Pn that accounts for carboxylation and CO diffusion processes, and assumes that the ratio of internal conductance to g(s) is constant, was constructed, and its goodness of fit was examined. KEY RESULTS: Considerable genotypic differences in Pn were evident for rice throughout development in both the pot and field experiments. The genotypic variation of Pn was correlated with that of g(s) at a given stage, and the change of Pn with plant development was closely related to the change of N. The variation of g(s) among genotypes was independent of that of N. The model explained well the variation in Pn of the ten genotypes grown under different conditions at different developmental stages. Conclusions The response of Pn to increased N differs with g(s), and the increase in Pn of genotypes with low g(s) is smaller than that of genotypes with high g(s). Therefore, simultaneous improvements of these two traits are essential for an effective breeding of rice genotypes with increased Pn.     

Spatial distribution of leaf water-use efficiency and carbon isotope discrimination within an isolated tree crown

The spatial variations in the stable carbon isotope composition (δ¹³C) of air and leaves (total matter and soluble sugars) were quantified within the crown of a well‐watered, 20‐year‐old walnut tree growing in a low‐density orchard. The observed leaf carbon isotope discrimination (Δ) was compared with that computed by a three‐dimensional model simulating the intracanopy distribution of irradiance, transpiration and photosynthesis (previously parameterized and tested for the same tree canopy) coupled to a biophysically based model of carbon isotope discrimination. The importance of discrimination associated with CO₂ gradients encountered from the substomatal sites to the carboxylation sites was evaluated. We also assessed by simulation the effect of current irradiance on leaf gas exchange and the effect of long‐term acclimation of photosynthetic capacity and stomatal and internal conductances to light regime on intracanopy gradients in Δ. The main conclusions of this study are: (i) leaf Δ can exhibit important variations (5 and 8‰ in total leaf material and soluble sugars, respectively) along light gradients within the foliage of an isolated tree; (ii) internal conductance must be taken into account to adequately predict leaf Δ, and (iii) the spatial variations in Δ and water‐use efficiency resulted from the short‐term response of leaf gas exchange to variations in local irradiance and, to a much lesser extent, from the long‐term acclimation of leaf characteristics to the local light regime.