羊草气孔导度的Jarvis-类模型

在干旱半干旱气候条件下,土壤水分状况通常是决定植物气孔导度的重要因素,现有气孔导度模型Jarvis-类和耦合模型(或光合-导度模型)未充分考虑这一因素对气孔导度的影响。本文以Jarvis气孔导度模型为基础,提出一个充分考虑土壤水分状况因素的气孔导度模型。该模型对羊草连续两年(1998~1999)野外实地观测结果拟合良好(R2=0.603),预测能力较线性回归方程(R2=0.361)有明显提高。     

冬小麦叶片气孔导度模型水分响应函数的参数化

植物气孔导度模型的水分响应函数用来模拟水分胁迫对气孔导度的影响过程, 是模拟缺水环境下植物与大气间水、碳交换过程的关键算法。水分响应函数包括空气湿度响应函数和土壤湿度(或植物水势)响应函数, 该研究基于田间实验观测, 分析了冬小麦(Triticum aestivum)叶片气孔导度对不同空气饱和差和不同土壤体积含水量或叶水势的响应规律。一个土壤水分梯度的田间处理在中国科学院禹城综合试验站实施, 不同水分胁迫下的冬小麦叶片气体交换过程和气孔导度以及其他的温湿度数据被观测, 同时观测了土壤含水量和叶水势。实验数据表明, 冬小麦叶片气孔导度对空气饱和差的响应呈现双曲线规律, 变化趋势显示大约1 kPa空气饱和差是一个有用的阈值, 在小于1 kPa时, 冬小麦气孔导度对空气饱和差变化反应敏感, 而大于1 kPa后则反应缓慢; 分析土壤体积含水量与中午叶片气孔导度的关系发现, 中午叶片气孔导度随土壤含水量增加大致呈现线性增加趋势, 但在平均土壤体积含水量大于大约25%以后, 气孔导度不再明显增加, 而是维持在较高导度值上下波动; 冬小麦中午叶片水势与相应的气孔导度之间, 随着叶水势的增加, 气孔导度呈现增加趋势。根据冬小麦气孔导度对空气湿度、土壤湿度和叶水势的响应规律, 研究分别采用双曲线和幂指数形式拟合了水汽响应函数, 用三段线性方程拟合了土壤湿度响应函数和植物水势响应函数, 得到的参数可以为模型模拟冬小麦的各类水、热、碳交换过程采用。     

羊草叶片气孔导度对环境因子的响应模拟

准确定量描述植物气孔对环境的响应是了解植物光合作用机理、预测植物生产力及其大气－植被－土壤系统中水分和热量交换的关键。利用松嫩平原盐碱化草地羊草光合生理特征的野外观测数据,分析了羊草叶片气孔导度对环境因子的反应,结果表明：羊草叶片气孔导度对环境因子变化敏感,尤其对瞬时光合有效辐射（ＰＡＲ）、叶片与空气间的水汽压亏损（ＶＰＤ）和空气温度（Ｔａ）反应十分明显。依据野外实测资料对国际上两类代表性气孔导度     

冬小麦群体叶片气孔导度差异性分析

对冬小麦 (Triticum aestivum L.)两个品种 3个生育期测定和分析了群体叶片的气孔导度。结果表明 ,近轴面气孔导度比远轴面的大 ,二者之比呈近似正态分布 ,最大概率出现约为 1 .5,该比值在冠层垂直方向变化不显著 ,但随季节而变 ,抽穗期最大。叶片气孔导度从冠层顶部向下迅速递减 ,递减系数约为 0 .57,且各层日变化大致平行。冠层上部气孔活跃 ,导度日变化明显 ,下部日变化不大。同一叶片的气孔导度由近轴面的叶基 ,经由其叶中、叶尖 ,翻过尖点 ,再从远轴面的叶尖经叶中向叶基递减。对近轴面叶中、叶尖气孔导度与该面平均气孔导度相关系数较大 ,而对远轴面叶中、叶基气孔导度与该面气孔导度相关系数较大 ,据此建议测定近轴面气孔导度时 ,应以叶中、叶尖部位为最佳点 ,而测定远轴面气孔导度时 ,应以叶中、叶基部位为最佳点     

最优气孔行为理论和气孔导度模拟

气孔调节功能是陆地生态系统碳-水耦合过程中最重要的环节。与即时的气孔导度测量相比, 气孔导度斜率能有效地反映气孔导度对CO₂浓度、饱和水汽压亏缺和光合作用的敏感性, 包含了环境因子对光合作用和临界水分利用效率等的综合影响, 为研究全球变化下陆地生态系统碳-水耦合关系提供了一个简明且综合的框架。气孔导度模型从经验模型、半经验模型发展到机理模型, 经过很多学者的改进, 但是模型参数的生物学意义和变化规律还不明确。鉴于气孔导度斜率方面研究的重要性和研究的不足, 为了加强对气孔导度调节规律的认识, 并减少气孔导度模拟的不确定性, 该文主要综述了长期以来国内外关于最优气孔行为理论和气孔导度模拟方面的研究成果, 其中包括广泛使用的气孔导度模型及参数意义, 讨论影响气孔导度斜率的主要因素以及气孔导度机理模型的应用, 并对最优气孔行为理论和气孔导度模拟方面的研究做了简单展望。     

羊草叶片气孔导度特征及数值模拟

对松嫩平原草地羊草叶片气孔导特征及与环境因子关系的研究结果表明,羊草叶片气孔导度日变化与环境因子密切相关,晴天表现为双峰曲线,阴天为单峰曲线,同时叶片气孔导度（gs)对瞬时光合有效辐射（PAR）,叶片与空气间的水汽压亏损（VPD）,空气温度（Ta）反应十分明显,依据野外实测资料,在对国际上两类代表性气孔导度模型验证比较的基础上,建立了适用于羊草草原的羊草叶片气孔导度对环境因子的响应模型gs=PAR(2.01Ta^2 147.74Ta-2321.11)/(444.62 PAR)(-538.04 VPD).     

植物气孔导度的机理模型

Ball-Berry气孔导度模型及其修正模型是评价植物叶片气孔调节的重要工具。该文从CO₂分子在叶片气孔中扩散这个最基本的物理过程出发, 应用物理学中的分子扩散和碰撞理论、流体力学与植物生理学等知识, 严格推导出叶片气孔导度的机理模型。利用美国Li-Cor公司生产的Li-6400光合作用测定仪控制CO₂浓度、湿度和温度, 测量了华北平原冬小麦(Triticum aestivum)的光响应数据和气孔导度数据。拟合结果表明: 推导的气孔导度机理模型较之Ball-Berry气孔导度模型和Tuzet等气孔导度模型, 能更好地描述冬小麦的气孔导度与净光合速率之间的关系。如果用气孔导度的机理模型耦合光合作用对光响应的修正模型, 则耦合模型可以很好地描述华北平原冬小麦叶片气孔导度对光强的响应曲线, 并可直接估算冬小麦的最大气孔导度和对应的饱和光强, 同时可以研究最大气孔导度是否与最大净光合速率同步的问题。拟合结果还表明: 冬小麦在30 ℃、560 μmol·mol^-1CO₂, 或在32 ℃、370 μmol·mol^-1CO₂条件下, 最大气孔导度与最大净光合速率并不同步。     

胡杨叶片气孔导度特征及其对环境因子的响应

依据2005年对极端干旱区荒漠河岸林胡杨的观测资料,对胡杨气孔运动进行了分析研究以揭示胡杨的水分利用特征与抗旱机理。结果表明:(1)胡杨叶片气孔导度日变化呈现为周期波动曲线,其波动周期为2 h,傍晚(20:00)波动消失;净光合速率和蒸腾速率与气孔导度的波动相对应而呈现同步周期波动。(2)胡杨的阳生叶气孔导度高于阴生叶,且不同季节气孔导度值不同,阳生叶气孔导度的季节变幅大于阴生叶。(3)胡杨气孔导度与气温、相对湿度和叶水势有显著相关关系,当CO2浓度较小时,胡杨气孔导度随CO2浓度的增加而增加,当CO2浓度达到一定值后气孔导度不再增加,反而随CO2浓度的增加大幅度降低。(4)胡杨适应极端干旱区生境的气孔调节机制为反馈式反应,即由于叶水势降低导致气孔导度减小,从而减少蒸腾耗水,达到节约用水、适应干旱的目的,表明胡杨的水分利用效率随气孔限制值的增大而减小,二者呈显著负相关。     

额济纳荒漠河岸胡杨林叶片气孔导度与微环境因子关系的模拟研究

以额济纳荒漠河岸胡杨(Populus euphratica)为研究对象,利用LI-6400光合测定仪于2005年5~9月份观测了胡杨叶片气体交换数据,研究了胡杨叶片气孔导度与光合速率、光合有效辐射与光合速率之间的关系.结果表明:(1)胡杨叶片净光合速率随气孔导度的增大而升高,但当气孔导度增加到一定值后,光合速率的增加变缓慢直至平稳,并主要是非气孔限制因素造成的;Ball-Berry模型(B-B模型)能够很好地描述气孔导度与光合速率之间的关系(R2=0.92).(2)叶片净光合速率随着有效辐射的变化符合非直角双曲线规律(R2=0.99).(3)B-B模型和非直角双曲线光合模型耦合后模拟值与观测值之间存在很好的正相关性(r=0.93),但耦合模型的模拟值还是较实测值偏大.因此,在干旱区还必须考虑水分限制因素对气孔开闭的控制作用,进一步构建适合干旱区生态系统特点的水-碳耦合循环机理模型.     

干旱条件下ABA与气孔导度和叶片生长的关系

介绍了干旱情况下ＡＢＡ的产生、运输,以及根源ＡＢＡ与气孔导度和叶片生长之间的定性、定量关系,并对干旱时叶片ＡＢＡ的重新分布,以及引起气孔关闭和抑制生长的机理作了叙述。     

A reinterpretation of stomatal responses to humidity

The stomatal conductance (g) for single leaves and the equivalent canopy conductance for stands of vegetation are often represented in models as empirical functions of saturation vapour pressure deficit or relative humidity. The mechanistic basis of this dependence is very weak. A reanalysis of 52 sets of measurements on 16 species supports the conclusion of Mott & Parkhurst (1991, Plant, Cell and Environment 14, 509–515) that stomata respond to the rate of transpiration (E) rather than to humidity per se. In general, ?g/?E is negative and constant so that the relation between g and E can be defined by two parameters: a maximum conductance g_m obtained by extrapolation to zero transpiration, and a maximum rate of transpiration E_m obtained by extrapolation to zero conductance. Both parameters are shown to be functions of temperature, CO₂ concentration, and soil water content. Exceptionally, transpiration rate and conductance may decrease together in very dry air, possibly because of patchy closure of stomata.     

Interpretation of an empirical model for stomatal conductance in terms of guard cell function

An empirical model for stomatal conductance (g), proposed by Leuning (1995, this issue) as a modification of Ball, Woodrow & Berry's (1987) model, is interpreted in terms of a simple, steady-state model of guard cell function. In this model, stomatal aperture is a function of the relative turgor between guard cells and epidermal cells. The correlation between g and leaf surface vapour pressure deficit in Leuning's model is interpreted in terms of stomatal sensing of the transpiration rate, via changes in the gradient of total water potential between guard cells and epidermal cells. The correlation between g, CO₂ assimilation rate and leaf surface CO₂ concentration in Leuning's model is interpreted as a relationship between the corresponding osmotic gradient, irradiance, temperature, intercellular CO₂ concentration and stomatal aperture itself. The explicit relationship between osmotic gradient and stomatal aperture (possibly describing the effect of changes in guard cell volume on the membrane permeability for ion transport) results in a decrease in the transpiration rate in sufficiently dry air. Possible extension of the guard cell model to include stomatal responses to soil water status is discussed.     

Ontogenetic changes in stomatal size and conductance of sunflowers

The ontogenetic changes in stomatal size, frequency and conductance (g_s) on abaxial and adaxial leaf surfaces of sunflower plants (Helianthus annuus L. Russian Mammoth) were examined under controlled environmental conditions. The stomatal frequency on the adaxial and abaxial leaf surfaces decreased with leaf ontogeny and insertion level. The ratio of adaxial to abaxial stomatal frequency did not change with leaf ontogeny and insertion level, and 42–44% of total stomata was apportioned to the adaxial surface. Ontogenetic changes in stomatal pore length were detected and increased with ontogenesis. The stomatal length of both leaf surfaces had linear relationships with leaf area. Ontogenetic changes in g_s were similar between the two surfaces. However the adaxial g_s was lower than abaxial g_s in leaves of higher insertion levels. Conductance had a linear relationship with width x frequency but not with pore area.     

The effect of tree height on crown level stomatal conductance

Variation in stomatal conductance is typically explained in relation to environmental conditions. However, tree height may also contribute to the variability in mean stomatal conductance. Mean canopy stomatal conductance of individual tree crowns (G_Si) was estimated using sap flux measurements in Fagus sylvatica L., and the hypothesis that G_Si decreases with tree height was tested. Over 13 d of the growing season during which soil moisture was not limiting, G_Si decreased linearly with the natural logarithm of vapour pressure deficit (D), and increased exponentially to saturation with photosynthetic photon flux density (Q_o). Under conditions of D = 1 kPa and saturating Q_o, G_Si decreased by approximately 60% with 30 m increase in tree height. Over the same range in height, sapwood‐to‐leaf area ratio (A_S:A_L) doubled. A simple hydraulic model explained the variation in G_Si based on an inverse relationship with height, and a linear relationship with A_S:A_L. Thus, in F. sylvatica, adjustments in A_S:A_L partially compensate for the negative effect of increased flow‐path length on leaf conductance. Furthermore, because stomata with low conductance are less sensitive to D, gas exchange of tall trees is reduced less by high D. Despite these compensations, decreasing hydraulic conductance with tree height in F. sylvatica reduces carbon uptake through a corresponding decrease in stomatal conductance.     

Effects of humidity on short-term responses of stomatal conductance to an increase in carbon dioxide concentration

The magnitude of the response of stomatal conductance to a change in the concentration of carbon dioxide external to the leaf from 350 to 700 cm³ m^–3 was found to be extremely variable from day to day in the field in Glycine max , Hordeum vulgare and Triticum aestivum . It was found that the leaf-to-air water vapour pressure difference (LAVPD) during the midday measurements of the stomatal response to carbon dioxide affected the magnitude of the response. On days when LAVPD was low, no significant change in conductance occurred with the increase in carbon dioxide concentration. When LAVPD was higher, conductance decreased by 24–52% with the increase in carbon dioxide within a few minutes. The sensitivity of conductance was approximately linearly related to LAVPD in wheat and barley. Experiments with G. max in the field indicated that, on days with low LAVPD, increasing the LAVPD just around the measured portion of a leaflet made stomatal conductance responsive to increased carbon dioxide. This result was also obtained under laboratory conditions with G. max , Helianthus annuus and Amaranthus retroflexus . In G. max , it was determined that leaves in which conductance was not responsive to the increase in carbon dioxide could be made responsive even at low LAVPD by the injection of abscisic acid into their petioles. Because it is known that abscisic acid sensitizes stomata to carbon dioxide, these results are consistent with the idea that abscisic acid may be involved in the response of stomatal conductance to changes in LAVPD.     

Inheritance of stomatal conductance in cotton (Gossypium barbadense)

Stomatal conductance in improved Pima cotton cultivars (Gossypium barbadense) has been previously shown to be positively associated with heat resistance and yield potential. In the present study we determined the mode of inheritance of stomatal conductance in crosses of six G. barbadense parents varying in origin, degree of agronomic development and stomatal conductance. Parents included a primitive tropical cotton (B368), two obsolete cultivars (St Vincent V135, Pima 32), one modern commercial line (Pima S-6) and two elite genotypes of the Pima germplasm (P70, P73). These lines showed distinct differences in stomatal conductance, under greenhouse and field conditions. The primitive B368 had the lowest conductance, and the elite lines the highest. Generation means analysis was used to quantify genetic effects in the crosses P70 × St Vincent V135, Pima S-6 × B368, Pima S-6 × Pima 32, P73 × Pima 32 and P73 × Pima S-6. Best-fit models of the inheritance of stomatal conductance varied in complexity from a simple additive-dominance model in the cross P70 × St. Vincent V135 to models displaying digenic epistatic interactions in the remaining crosses. Significant additive mean effects for stomatal conductance were detected in all crosses. Dominance mean effects were significant in the crosses P73 × Pima 32 and P73 × Pima S-6. Broadsense heritability estimates of stomatal conductance were relatively low (0.16–0.44) in all crosses except Pima S-6 × B368 (0.74). Results also show that the mode of inheritance of stomatal conductance is multigenic, and may have maternal as well as nuclear components. Recouping higher stomatal conductance levels from genetically wider crosses appears feasible and could proceed at a moderate rate. Fixing higher levels of stomatal conductance in populations from crosses of elite germplasm may be more difficult because of the presence of dominant mean effects and digenic epistatic interactions.     

Effects of patchy stomatal closure on gas exchange measurements following abscisic acid treatment

Gas exchange data and images of leaf fluorescence were collected concurrently as stomata responded to abscisic acid (ABA) application. When 10^?5kmolm^?3 ABA was applied to the transpiration stream in a short pulse, stomatal conductance (g_s), photosynthesis (A) and intercellular CO2 concentration (C_i) decreased rapidly after a short lag period and became approximately constant after 2h. There was an apparent reduction in the A versus c1 relationship as stomata closed, but the data returned to the A versus C1 curve while stomatal conductance was constant or slowly rising during the second hour after ABA treatment. Larger amounts of ABA administered during the pulse caused larger deviations from the A versus c1 relationship. When 10^?7kmolm^?3 ABA was applied continuously through the transpiration stream, gs, A and Cⁱ decreased, but there was no substantial deviation from the A versus c{ curve. Fluorescence images were patchy as stomata closed for all experiments, but became slowly more uniform during the time that gas exchange was returning to the A versus Cj curve. The distribution of con-ductance among patches was not bimodal, and larger devi-ations from the A versus ct curve had greater ranges of pixel values and more pixel values representing low values of Cj during stomatal closure than did experiments show-ing small or no deviation. Estimates of A and gs from fluo-rescence images compared favourably with measured val-ues in most cases, suggesting that the patchy distributions of fluorescence were caused by patchy distributions of stomatal conductance and that apparent reductions in the A versus ct relationship were the result of these patchy stomatai distributions and not direct effects of ABA on mesophyll functioning. The data show that stomatal patches can be temporary and that patchiness may not be reflected in gas exchange data if the range of stomatal con-ductances is not large. These observations may explain some of the discrepancies among previous studies concerning the effect of ABA on the A versus Cⁱ relationship.     

Hydraulic conductance, stomatal conductance, and maximal photosynthetic rate in bean leaves

A positive correlation was found between steady state values of hydraulic (L _pA) and stomatal conductance (g _s) of French bean leaves: both were lower in the dark than in the light and lower in water-deficient plants than in the well-watered ones. The relative rate of stomatal opening after a pressure rise in the xylem was also positively related to L _pA. The L _pA and g _s were both related to the maximal photosynthetic rate at saturating CO2 concentrations. This revised version was published online in June 2006 with corrections to the Cover Date.