麦田冠层气孔导度的分层研究

小麦灌浆期和乳熟期冠层各层叶片上、下表面的气孔导度之间呈正相关关系;冠层不同层的叶片气孔导度从早到傍晚有平行变化的趋势,数值上存在较大的差异,一般从冠层上到下递减。经分析,这主要与冠层叶片接受的光强自上而下递减有关,且这时所对应的叶片水势自冠层上到下递增的幅度大。测算结果表明,冠层气孔导度白天亦呈明显的日变化,灌浆期的值大于乳熟期的值。     

叶片生长进程中气孔发育对叶温调节的影响

鉴于气孔发育影响气孔导度和蒸腾速率,推测气孔发育可能影响叶温调节。为验证这一假设并阐述相关规律,在控光和控温条件下研究了冬青卫矛和华北紫丁香气孔发育、气孔导度、蒸腾速率及其与叶温的关系。结果表明,伴随冬青卫矛、华北紫丁香叶片生长气孔逐渐增大,但气孔密度下降;在此过程中,气孔导度和蒸腾速率逐步提高,而叶片温度降低;尽管冬青卫矛和华北紫丁香叶片的气孔密度和大小差异很小,但华北紫丁香近轴侧和远轴侧均有气孔分布,而冬青卫矛则只有远轴侧分布气孔,且相同条件下华北紫丁香的气孔导度和蒸腾速率高、叶温低。因此,气孔发育能够促进气孔导度和蒸腾速率提高,有助于降低叶温;近轴侧气孔可能更有利于蒸腾降温。     

田间小麦群体内叶片气孔阻力垂直差异研究

根据田间实测资料, 本文给出了田间小麦群体中叶片不同部位和冠层不同层次叶面气孔阻力的差异, 及其与环境因素特别是土壤水分含量关系的分析结果。结果表明, 小麦冠层各不同层次和叶片不同部位的气孔阻力有明显差异。就冠层上部单个叶片而言, 由叶尖到叶基气孔阻力逐渐增大;叶片正面的气孔阻力小于叶片反面对应部位的气孔阻力。对于水汽和CO₂传导的贡献70%是由冠层上部的两片叶子给出的;群体内不同层次叶片相应部位的气孔阻力是由冠层上部向下逐渐增大, 冠层不同层次叶片和叶片不同部位的气孔对环境因素变化的反空不同, 如当土壤水分不足时, 群体下部叶片和叶片反面的气孔张度首先变小, 且变小的程度较大。     

植物气孔导度的环境响应模拟及其尺度扩展

气孔导度是衡量植物和大气间水分、能量及CO2平衡和循环的重要指标,探讨气孔导度与环境因子的关系及其模拟,以及气孔导度在叶片、冠层及区域尺度间的尺度转换及累积效应,对更好地认识植被与大气间的水热运移过程,合理评价植被在陆面过程中的地位和作用都具有重要意义。从植物气孔导度与环境因子的关系、气孔导度模拟以及尺度扩展三个方面,对前人的研究成果进行了概括总结。从叶片和冠层两个尺度出发,归纳总结了前人对于不同植物气孔导度与环境因子关系的研究成果,发现由于不同植物的遗传特性、测定时的环境、时间尺度的不同,以及未考虑各个环境因子的相互作用对气孔导度的影响,由此得到的气孔导度与环境因子之间的关系也不尽一致。对各单一环境因子与气孔导度的关系,给出了生理学解释,从根本上说明了环境因子变化对气孔导度的影响,而研究环境因子对气孔导度的综合影响时,应对各环境因子进行系统控制与同步观测。模拟计算植物气孔导度的模型主要有Jarvis模型和BWB模型两类,这些模型的模拟能力随着研究对象、试验区域、环境条件的改变而存在一定的差异,在具体使用时应结合实际情况选择最优模型进行模拟。除上述常用模型外,还总结了其他学者分别从不同角度提出的新的模型,对现有气孔导度模型进行了全面的总结。从叶片-冠层、冠层-区域两个方面归纳总结了前人关于气孔导度尺度扩展的研究成果,发现叶片-冠层的尺度扩展研究较成熟而冠层-区域的尺度扩展在模拟精度的验证方面存在困难。针对以下几个方面提出了今后气孔导度的研究重点:(1)结合研究对象所在的区域及环境条件,选择最优模型进行模拟;(2)综合考虑环境因子之间的相互作用及其对气孔导度的累积影响;(3)BWB模型与光合模型的耦合;(4)提高大尺度范围内的气孔导度模拟精度。     

七种热带雨林树苗叶片气孔特征及其可塑性对不同光照强度的响应

对生长在荫棚3种不同光照条件下和全自然光下的热带雨林4个冠层种(望天树、绒毛番龙眼、团花、红厚壳)和3个中层种(玉蕊、藤黄、滇南风吹楠)树苗叶片气孔特征以及它们的可塑性进行了研究、结果表明,这些植物的气孔全部着生在远轴面．7种植物中,玉蕊和绒毛番龙眼的气孔密度较大,滇南红厚壳和团花的保卫细胞最长．随光强的增大,气孔密度和气孔指数增大,单位叶气孔数在低光强下较大．除团花外,其它植物叶片气孔导度在50％光强处最大,而光强对保卫细胞的长度影响不显著．相关分析表明,气孔密度与植物单位叶的面积呈负相关。而与气孔导度的相关性不显著、尽管两种不同生活型植物气孔指数和单位叶气孔数在不同光强下的可塑性差异较小,但冠层树种气孔密度和气孔导度的可塑性显著高于中层树种．     

水稻叶片气孔导度与冠层反射光谱的定量关系分析

研究了不同土壤水氮条件下水稻(Oryza sativa)叶片气孔导度与冠层光谱反射特征的量化关系。结果表明,不同水分处理下,水稻不同叶位气孔导度变化趋势为:GsL1>GsL2>GsL3>GsL4。高于W3水分条件下,高氮处理的叶片气孔导度高于低氮处理,而低于W3水分条件下,高低氮处理条件下叶片气孔导度差异不显著。发现比值指数R(1 650,760)与不同叶位叶片及不同层次叶片气孔导度的相关性大小为:GsL1>GsL12>GsL123>GsL1234>GsL2>GsL3>GsL4(水稻顶部自上而下第一、二、三、四叶以及自上而下顶部2张、3张、4张叶片的气孔导度值分别表示为:GsL1、GsL2、GsL3、GsL4、GsL12、GsL123和GsL1234),而顶1叶气孔导度与叶面积指数的乘积(冠层叶片气孔导度)同比值指数R(1 650,760)相关程度更高。R(1 650,760)与顶1叶和冠层叶片气孔导度之间皆呈极显著的幂函数负相关。利用不同年份的不同水稻试验对两者的监测模型进行了检验,模型的检验误差RMSE分别为0.05和0.24,表明比值指数R(1 650,760)可以较好地监测不同水氮条件下水稻叶片的气孔开闭特征。     

华北山区侧柏冠层气孔导度特征及其对环境因子的响应

冠层气孔导度(g_s)是衡量冠层-大气界面水汽通量的重要生物学常数,研究其特征及对环境因子的响应,能为开展森林冠层水汽交换过程的机理性研究提供理论依据.于2014年利用SF-L热扩散式探针测定了侧柏的树干液流密度(J_s),同步监测光合有效辐射(PAR)、饱和水汽压差(VPD)、气温(T)等环境因子,计算侧柏的冠层气孔导度特征并分析其对各环境因子的响应.结果表明: 侧柏液流密度的日变化总体呈双峰曲线,生长季高于非生长季,且胸径越大液流密度越大;冠层气孔导度日变化与单位叶面积冠层蒸腾(E_L)趋势相近,均呈双峰曲线,生长季的冠层气孔导度和蒸腾较非生长季略高.侧柏冠层气孔导度与空气温度呈抛物线关系,在10 ℃左右冠层气孔导度达到峰谷;光合有效辐射以400 μmol·m^-2·s^-1为界,小于该阈值两者呈正相关关系,大于该阈值则冠层气孔导度受其影响较小;与饱和水汽压差呈负对数函数关系,随饱和水汽压差增大而逐渐降低.较高的空气温度和光合有效辐射、较低的饱和水汽压差有利于侧柏形成较大的冠层气孔导度,进而促进冠层蒸腾.     

植被-大气相互作用中的气孔导度及其尺度转换

气孔导度是衡量植物和大气间水分、能量及CO2平衡和循环的重要指标,探讨气孔导度在叶片、冠层及区域尺度间的尺度转换及累积效应,对更好地认识植被与大气间的水热运移过程,合理评价植被在陆面过程中的地位和作用具有重要意义.本文着重从叶片尺度气孔导度模拟、气孔导度在冠层尺度的累积表现、冠层到区域尺度转换研究及气孔导度累积效应在陆面过程模型中的作用等4个层次总结了近期国内外研究状况,指出其中存在的异质性等问题,并就今后应加强多尺度间的同步观测提出了展望.     

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     

Variation of Leaf Stomatal Conductance in Winter Wheat Canopy

Leaf stomatal conductance measured and analysed in the canopies of two winter wheat varieties in the field revealed that the probability of adaxial to abaxial conductance ratio followed an approximately normal distribution with a peake value of about 1.5. The ratio changed with the developmental stages being maximium at the heading stage. Leaf stomata in wheat of the upper part of the canopy were more active and showed more pronounced diurnal change of conductance than those of the lower part. Stomatal conductance decreased from top to bottom in canopy as a negative exponential function. By comparing adaxial and abaxial conductances in the apical, middle and basal parts of a leaf, the distribution of the stomatal conductances of a wheat leaf was as follows: a steady decrease from the basal part of adaxial, through the middle and apical parts of the adaxial surface turning to the apical part of abaxial, and then the middle and lastly, the basal part of abaxial. Based on values of the correlation coefficients among the various stomatal conductance and average stomatal conductance, the authors suggested that optimal apical measurement of stomatal conductance would be at the middle and apical parts and that of abaxial would be at middle and basal parts.     

Asymmetric responses of adaxial and abaxial stomata to elevated CO2: impacts on the control of gas exchange by leaves

The response of adaxial and abaxial stomatal conductance in Rumex obtusifolius to growth at elevated atmospheric concentrations of CO₂ (250 μmol mol^?1 above ambient) was investigated over two growing seasons. The conductance of both the adaxial and abaxial leaf surfaces was found to be reduced by elevated concentrations of CO₂. Elevated CO₂ caused a much greater reduction in conductance for the adaxial surface than for the abaxial surface. The absence of effects upon stomatal density indicated that the reductions were probably the result of changes in stomatal aperture. Partitioning of gas exchange between the leaf surfaces revealed that increased concentrations of CO₂ caused increased rates of photosynthesis only via the abaxial surface. Additionally, leaf thickness was found to increase during growth at elevated concentrations of CO₂. The tendency for these amphistomatous leaves to develop a distribution of conductance approaching that of hypostomatous leaves clearly reduced their maximum photosynthetic potential. This conclusion was supported by measurements of stomatal limitation, which showed greater values for the adaxial surfaces, and greater values at elevated CO₂. This reduction in photosynthesis may in part be caused by higher diffusive limitations imposed because of increased leaf thickness. In an uncoupled canopy, asymmetrical stomatal responses of the kind identified here may appreciably reduce transpiration. Species which show symmetrical responses are less likely to show reduced transpirational rates, and a redistribution of water loss between species may occur. The implications of asymmetrical stomatal responses for photosynthesis and canopy transpiration are discussed.     

Abaxial and adaxial stomata from Pima cotton (Gossypium barbadense L.) differ in their pigment content and sensitivity to light quality

Sensitivity to light quality and pigment composition were analysed and compared in abaxial and adaxial stomata of Gossypium barbadense L. (Pima cotton). In most plants, abaxial (lower) stomatal conductances are higher than adaxial (upper) ones, and stomatal opening is more sensitive to blue light than to red. In greenhouse-grown Pima cotton, abaxial stomatal conductances were two to three times higher than adaxial ones. In contrast, adaxial stomatal conductances were 1·5 to two times higher than abaxial ones in leaves from growth chamber-grown plants. To establish whether light quality was a factor in the regulation of the relationship between abaxial and adaxial stomatal conductances, growth-chamber-grown plants were exposed to solar radiation outdoors and to increased red light in the growth chamber. In both cases, the ratios of adaxial to abaxial stomatal conductance reverted to those typical of greenhouse plants. We investigated the hypothesis that adaxial stomata are more sensitive to blue light and abaxial stomata are more sensitive to red light. Measurements of stomatal apertures in mechanically isolated epidermal peels from growth chamber and greenhouse plants showed that adaxial stomata opened more under blue light than under red light, while abaxial stomata had the opposite response. Using HPLC, we quantified the chlorophylls and carotenoids extracted from isolated adaxial and abaxial guard cells. All pigments analysed were more abundant in the adaxial than in the abaxial guard cells. Antheraxanthin and β-carotene contents were 2·3 times higher in adaxial than in abaxial guard cells, comparing with ad/ab ratios of 1·5–1·9 for the other pigments. We conclude that adaxial and abaxial stomata from Pima cotton have a differential sensitivity to light quality and their distinct responses are correlated with different pigment content.     

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.     

Effects of low level O3 exposure on leaf diffusive conductance and water-use efficiency in hybrid poplar

Abstract Young, amphistomatous hybrid poplar (Populus deltoides x trichocarpa) plants were exposed daily to either background (0.025 cm³ m^-3) or elevated (0.125 cm³ m^-3) concentrations of O₃. Levels of abaxial and adaxial leaf conductance were affected interactively by pollutant treatment, leaf age, and photon fluence rate. Consequently, conductance in O₃-treated leaves was sometimes higher and sometimes lower than in comparable control leaves, depending on leaf age or level of photon fluence rate. For example, at low photon fluence rate or in the dark, conductance was greater in O₃-treated than in control plants, while at high photon fluence rate that relationship was reversed. Exposure to O₃ also reduced the water-use efficiency and range of leaf conductance of individual leaves, and altered the relationship between the conductances of the two leaf surfaces (the ratio of abaxial to adaxial leaf conductance was increased). Furthermore, O₃ treatment resulted in diminished stomatal control of water loss; excised O₃-treated leaves had higher conductances and wilted sooner than excised control leaves of identical ages. Overall, the data indicate that exposure to O₃ resulted in impaired stomatal function.     

小麦和玉米叶片两面气孔导性的差异及其与环境因素的关系

气孔是作物与大气进行水汽和CO_2交换的通道,当环境条件发生变化时,作物通过调节气孔的开度来控制叶片内部和外部的气体交换速率。作物群体中叶片两面的小生境有明显差异,在长期进化和适应中形成了两面气孔特性     

Stomatal characteristics,conductance ratios,and drought-induced leaf modifications of semiarid grassland species

Seventeen greenhouse-grown grasses from the Nebraska Sandhills region were surveyed for foliar stomatal density and distribution, closed guard cell lengths, open stomatal apertures, and surface characteristics (using scanning electron microscopy), surface conductance (using a steady-state porometer), and drought-induced leaf modifications. Leaves of C₃ species exhibited a proclivity toward being amphistomatic or hyperstomatic, while C₄ species tended to be more hypostomatic. Leaf modification, when it occurred, resulted in the enshrouding of the adaxial surface. Conductance data showed functional amphistomaty in most species, revealing differential functioning of adaxial and abaxial stomata. Conductance patterns were not closely related to stomatal aperture per unit area leaf surface or to stomatal distribution patterns. Lowered adaxial: abaxial conductance ratios, increased stomatal density, reduced stomatal size, and less drought-induced leaf modification were seen in C₄ grasses as compared with C₃ grasses. C₃ range and C₃ meadow species did not differ in conductance ratios, density ratios, or stomatal size, although meadow species exhibited much greater drought-induced leaf modification. Postulations involving correlation of adaxial: abaxial conductance ratios to stomatal distribution patterns, and assumptions of stomatal distribution based upon habitat and/or photosynthetic pathway may be erroneous. These characteristics may be of limited usefulness as morphological indicators in the search for drought-tolerant ecotypes of prairie grasses.     

Abaxial and Adaxial Stomatal Density, Stomatal Conductances and Water Status of Bean Primary Leaves as Affected by Paclobutrazol

The plant growth retardant, paclobutrazol at 8.5 or 17.0 μM concentrations effectively inhibited the stem elongation and primary leaf expansion of bean seedlings. Although the retardant reduced the relative water content in well-watered plants, the water and pressure potentials remained high in the primary leaves. K⁺, Na⁺, Mg²⁺ and Ca²⁺ contents in the primary leaves of the paclobutrazol-treated plants were not significantly different from those in the control. The stomatal density increased on both surfaces but the length of guard cells was not reduced significantly on the adaxial epidermes of the paclobutrazol-treated primary leaves. The inhibitory effect of paclobutrazol on the abaxial stomatal conductances became more pronounced with time during the light period but the adaxial surfaces displayed similar or slightly higher conductances than those of the control. The transpiration rate on a unit area basis did not change significantly or increased in the treated leaves thus the reduced water loss of paclobutrazol-treated plants was due to the reduced leaf area. Stomatal conductances of the adaxial surfaces responded more intensively to exogenous abscisic acid and the total leaf conductance decreased faster with increasing ABA concentration in the control than in the paclobutrazol-treated leaves. Paclobutrazol, an effective inhibitor of phytosterol biosynthesis, not only amplified the stomatal differentiation but increased the differences between the adaxial and abaxial stomatal conductances of the primary leaves.