北京山区油松和元宝槭冠层气孔导度特征及其环境响应

叶片气孔是植物进行水汽交换的通道, 影响着植物的蒸腾和光合作用。然而叶片气孔行为受环境条件和树种类型的影响, 不同树种冠层气孔导度对环境因子响应的差异性, 以及在生长季不同时期叶片气孔对冠层蒸腾的调节作用是否会发生改变, 仍不清楚。该研究目的是通过探究各环境因子对不同树种冠层气孔导度的相对贡献率以及叶片气孔对冠层蒸腾的调节作用, 为深入了解植物水分利用状况和山区森林经营提供参考依据。于2018年生长季以北京八达岭国家森林公园内的58年生油松(Pinus tabuliformis)和39年生元宝槭(Acer truncatum)为研究对象, 利用热扩散技术对其树干液流进行连续监测, 并同步监测环境因子。利用彭曼公式计算冠层气孔导度(G_s)。主要结果: (1)油松和元宝槭日间G_s在日、月时间尺度上存在明显差异。5-7月油松和元宝槭日动态G_s均随饱和水汽压差(VPD)和太阳辐射(GR)的增加呈上升趋势, 上升持续时间比8月和9月长; 在月尺度上, 随着VPD、GR的降低和土壤湿度(VWC)的升高, G_s从5月到9月整体上升。(2)利用增强回归树法分析得到VWC和VPD对G_s的贡献率最大, 其次是GR、气温和风速。VWC和VPD对油松G_s的贡献率分别为66.4%和17.4%, 对元宝槭G_s的贡献率分别为54.8%和21.0%。(3)油松和元宝槭的dG_s/dlnVPD值与参考冠层气孔导度之间的斜率均显著高于0.6, 气孔调节作用相对较强。综上所述, 气孔对环境因子的响应在树种以及生长季不同时期之间存在差异, 为防止水分过度散失, 两树种在不同土壤水分条件下均通过严格的气孔调节控制蒸腾量。     

基部被子植物气孔性状与叶脉密度的关联进化

植物叶片通过气孔的水分蒸腾散失和叶脉的水分供应达到水分平衡,而基部被子植物在进化过程中叶片水分供应和蒸腾散失是否达到平衡或关联进化还缺乏了解。本研究以11种基部被子植物为材料,测定了气孔密度、气孔长度、叶脉密度和叶片厚度4个叶片性状,并结合系统发育树,利用系统发育独立对比的方法分析这些性状之间的关联进化。结果显示：沿进化方向,气孔密度和叶脉密度逐渐增加,而气孔长度和叶片厚度有减小的趋势;无论是否考虑系统发育的影响,气孔密度都与叶脉密度呈显著正相关关系,说明二者之间存在关联进化,并证实了基部被子植物叶片水分平衡假说;气孔密度和长度、叶脉密度均与叶片厚度呈显著线性相关,但在去除系统发育的影响后这种线性相关关系不再显著,说明叶片厚度与其它三个叶片性状不存在关联进化。本研究结果还表明,叶片的水分供应和散失乃至CO2通透性的平衡主导着基部被子植物叶片结构和功能的进化。     

Stomatal responses to non-local changes in PFD: evidence for long-distance hydraulic interactions

Interactions among stomata within a single areole have recently been reported, and evidence suggests that hydraulic mechanisms may be responsible for these interactions. Such interactions may play a role in patchy stomatal behaviour by coordinating stomatal behaviour within areoles. However, models suggest that longer‐distance interactions may be required to produce the large‐scale discoordination that is characteristic of stomatal patchiness. This study was undertaken to characterize long‐distance interactions between ‘artificial patches’ of stomata under varying conditions of evaporative demand and soil water stress. Gas‐exchange was monitored in two adjacent regions (‘patches’) of a wheat leaf by two independent gas mixing and analysis systems. When photon flux density (PFD) was changed in only one of these patches, stomatal conductance responded in both patches in a manner consistent with hydraulic interactions propagated by changes in xylem water potential. These data are discussed in the context of mechanisms for patchy stomatal conductance and implications for the design and analysis of gas‐exchange experiments.     

热带雨林木质藤本植物叶片性状及其关联

热带雨林中木质藤本植物较为丰富。随着全球气候变化加剧,木质藤本植物的丰富度具有不断增加的趋势,有可能对热带森林的结构、功能和动态产生重要影响。然而,目前对木质藤本响应环境变化的机制所知甚少。本研究以13个科20种热带雨林常见木质藤本植物为材料,测定了冠层叶片的17个形态特征及结构性状,并分析了性状间的相互关系。结果表明,叶片相对含水量的种间变异最小（变异系数为5%）,而上表皮厚度的种间变异最大（变异系数为80%）,其它性状的种间变异系数为24%~61%。木质藤本植物的叶脉密度、叶片密度均与气孔密度呈显著正相关,叶片干物质含量与比叶面积呈显著负相关。与相同生境的树木相比,木质藤本的叶面积更小、气孔密度和叶片密度更低、比叶面积更高,但两种植物类群的叶片横切面组织结构厚度无显著差异。研究结果对理解木质藤本植物的生态适应性具有重要意义。     

水稻OsYDA基因调控气孔发育的功能分析

气孔是植物特化的表皮结构,在植物蒸腾过程和与外界气体交换过程中起到重要作用。拟南芥YDA（AtYDA）是MAPK级联信号途径中的一种激酶（MAPKKK4）,它在叶片气孔的发育过程中起着负调控的作用。AtYDA功能缺失导致叶片气孔显著增加,而表达组成型激活形式的AtYDA（ΔN-YDA）则会导致表皮产生无气孔表型。本研究克隆了水稻中与AtYDA同源的2个基因OsYDA1和OsYDA2。在拟南芥中过量表达这2个基因都导致了叶片气孔密度的减少和叶片失水速率的降低。而表达ΔN-OsYDA1和ΔN-OsYDA2的转基因植株则呈气孔系数下降的表型。这表明OsYDA与AtYDA在调控气孔发育的功能上具有保守性。     

Transpiration and canopy conductance in a pristine broad-leaved forest of Nothofagus: an analysis of xylem sap flow and eddy correlation measurements

Summary Tree transpiration was determined by xylem sap flow and eddy correlation measurements in a temperate broad-leaved forest of Nothofagus in New Zealand (tree height: up to 36 m, one-sided leaf area index: 7). Measurements were carried out on a plot which had similar stem circumference and basal area per ground area as the stand. Plot sap flux density agreed with tree canopy transpiration rate determined by the difference between above-canopy eddy correlation and forest floor lysimeter evaporation measurements. Daily sap flux varied by an order of magnitude among trees (2 to 87 kg day^–1 tree^–1). Over 50% of plot sap flux density originated from 3 of 14 trees which emerged 2 to 5 m above the canopy. Maximum tree transpiration rate was significantly correlated with tree height, stem sapwood area, and stem circumference. Use of water stored in the trees was minimal. It is estimated that during growth and crown development, Nothofagus allocates about 0.06 m of circumference of main tree trunk or 0.01 m² of sapwood per kg of water transpired over one hour.Maximum total conductance for water vapour transfer (including canopy and aerodynamic conductance) of emergent trees, calculated from sap flux density and humidity measurements, was 9.5 mm s^–1 that is equivalent to 112 mmol m^–2 s^–1 at the scale of the leaf. Artificially illuminated shoots measured in the stand with gas exchange chambers had maximum stomatal conductances of 280 mmol m^–2 s^–1 at the top and 150 mmol m^–2 s^–1 at the bottom of the canopy. The difference between canopy and leaf-level measurements is discussed with respect to effects of transpiration on humidity within the canopy. Maximum total conductance was significantly correlated with leaf nitrogen content. Mean carbon isotope ratio was –27.76±0.27 (average ±s.e.) indicating a moist environment. The effects of interactions between the canopy and the atmosphere on forest water use dynamics are shown by a fourfold variation in coupling of the tree canopy air saturation deficit to that of the overhead atmosphere on a typical fine day due to changes in stomatal conductance.This paper is dedicated to Prof. Dr. O.L. Lange on the occasion of his 65th birthday     

不同水分条件下盆栽苹果树蒸腾速率动态模拟

土壤水分是制约作物产量和品质的主要环境因素之一,估算不同水分条件下的蒸腾速率（r3对于作物的优质高产和节水灌溉等具有十分重要的意义。通过构建蒸腾．气孔．光合耦合模型可模拟出不同水分条件下苹果树的蒸腾动态,模型参数根据逐步干旱条件下盆栽‘富士’苹果树试验获取。结果表明,Tr主要由饱合水气压差和气孔导度（G。）驱动,同时气象因子和土壤水势对其有强烈的交互作用影响。Tr随土壤水势的下降而减小,当土壤水势低于-0．4MPa时减小幅度更加显著。晴天时,G。在一天中呈双峰曲线,而一呈单峰曲线,最大值出现在13：00左右,约为3．6mmol·m-Ls。根据该模型可计算出不同水分条件下1株盆栽苹果树（总叶片积为O．26m-2）全天的蒸腾总量,供水充足时为652．1g,严重干旱时（土壤水势为-1．5MPa）为85．4g。实测值和模拟值的比较表明,该耦合模型能够模拟出不同土壤水分条件下盆栽苹果树的蒸腾动态以及土壤的含水量。     

Stomatal regulation by microclimate and tree water relations: interpreting ecophysiological field data with a hydraulic plant model

Dynamics in microclimate and physiological plant traits were studied for Pubescent oak and Scots pine in a dry inner-alpine valley in Switzerland, at a 10 min resolution for three consecutive years (2001-2003). As expected, stomata tended to close with increasing drought in air and soil. However, stomatal aperture in oak was smaller than in pine under relatively wet conditions, but larger under dry conditions. To explore underlying mechanisms, a model was applied that (i) quantifies water relations within trees from physical principles (mechanistic part) and (ii) assumes that signals from light, stomatal aperture, crown water potential, and tree water deficit in storage pools control stomata (systemic part). The stomata of pine showed a more sensitive response to increasing drought because both factors, the slowly changing tree water deficit and the rapidly changing crown water potential, closed the stomata. By contrast, the stomata of oak became less drought-sensitive as the closing signal of crown water potential was opposed by the opening signal of tree water deficit. Moreover, parameter optimization suggests that oak withdrew more water from the storage pools and reduced leaf water potentials to lower levels, without risking serious damage by cavitation. The new model thus suggests how the hydraulic water flow and storage system determines the responses in stomatal aperture and transpiration to drought at time scales ranging from hours to multiple years, and why pine and oak might differ in such responses. These differences explain why oaks are more efficient competitors during drought periods, although this was not the case in the extremely dry year 2003, which provoked massive leaf loss and, from July onwards, physiological activity almost ceased.     

Adjustments in hydraulic architecture of Pinus palustris maintain similar stomatal conductance in xeric and mesic habitats

We investigated relationships between whole-tree hydraulic architecture and stomatal conductance in Pinus palustris Mill. (longleaf pine) across habitats that differed in soil properties and habitat structure. Trees occupying a xeric habitat (characterized by sandy, well-drained soils, higher nitrogen availability and lower overstory tree density) were shorter in stature and had lower sapwood-to-leaf area ratio (A(S):A(L)) than trees in a mesic habitat. The soil-leaf water potential gradient (psiS - psiL) and leaf-specific hydraulic conductance (kL) were similar between sites, as was tissue-specific hydraulic conductivity (Ks) of roots. Leaf and canopy stomatal conductance (gs and Gs, respectively) were also similar between sites, and they tended to be somewhat higher at the xeric site during morning hours when vapour pressure deficit (D) was low. A hydraulic model incorporating tree height, A(S):A(L) and psiS-psiL accurately described the observed variation in individual tree G(Sref) (G(S) at D = 1 kPa) across sites and indicated that tree height was an important determinant of G(Sref) across sites. This, combined with a 42% higher root-to-leaf area ratio (A(R):A(L)) at the xeric site, suggests that xeric site trees are hydraulically well equipped to realize equal--and sometimes higher potential for conductance compared with trees on mesic sites. However, a slightly more sensitive stomatal closure response to increasing D observed in xeric site trees suggests that this potential for higher conductance may only be reached when D is low and when the capacity of the hydraulic system to supply water to foliage is not greatly challenged.     

树高对马占相思整树水分利用的效应

利用Granier热消散探针,于2004年观测了华南丘陵坡地常见绿化先锋树种马占相思(22年生)的树干液流,同时监测林冠上方的光合有效辐射、气温、相对湿度和0～30 cm的土壤体积含水量.结合树木的形态特征、液流密度和简化的Whitehead & Jarvis公式,分别计算了整树蒸腾、冠层气孔导度和叶面积/边材面积比值,分析了树高对整树蒸腾、冠层气孔导度和叶面积/边材面积比值的影响.结果表明：土壤水分充足时,马占相思整树蒸腾随树高呈二次多项式增加(P<0.01),冠层气孔导度日变化均呈“单峰”格型;在所有光合有效辐射范围内,高树的参比冠层气孔导度和冠层气孔导度对水汽压亏缺的敏感性均高于矮树;叶面积/边材面积比值为（1.837±0.048） m²·cm^-2,并与树高呈幂函数关系.随着树木高度的增加,马占相思没有发生明显的水力限制和补偿.     

Water use strategy affects avoidance of ozone stress by stomatal closure in Mediterranean trees—A modelling analysis

Both ozone (O₃) and drought can limit carbon fixation by forest trees. To cope with drought stress, plants have isohydric or anisohydric water use strategies. Ozone enters plant tissues through stomata. Therefore, stomatal closure can be interpreted as avoidance to O₃ stress. Here, we applied an optimization model of stomata involving water, CO₂, and O₃ flux to test whether isohydric and anisohydric strategies may affect avoidance of O₃ stress by stomatal closure in four Mediterranean tree species during drought. The data suggest that stomatal closure represents a response to avoid damage to the photosynthetic mechanisms under elevated O₃ depending on plant water use strategy. Under high-O₃ and well-watered conditions, isohydric species limited O₃ fluxes by stomatal closure, whereas anisohydric species activated a tolerance response and did not actively close stomata. Under both O₃ and drought stress, however, anisohydric species enhanced the capacity of avoidance by closing stomata to cope with the severe oxidative stress. In the late growing season, regardless of the water use strategy, the efficiency of O₃ stress avoidance decreased with leaf ageing. As a result, carbon assimilation rate was decreased by O₃ while stomata did not close enough to limit transpirational water losses.     

不同生境下入侵种假臭草叶片的气孔特征

【背景】探讨入侵种假臭草不同生境下气孔的变化规律,揭示假臭草种群在不同生境下所采取的生长对策及适应机制,可为入侵生物的防治提供参考。【方法】采取光学显微镜系统观察桉树林、木薯地、弃耕地、公路边4种生境下假臭草叶片的气孔特征。【结果】光照和土壤肥、水条件对假臭草叶片的气孔孔径（横轴方向和纵轴方向）、单个气孔器面积、气孔器总面积、气孔密度及气孔指数的影响显著。低光照及肥沃、湿润土壤生境与高光照及贫瘠、干旱土壤相比,假臭草的气孔孔径（横轴方向和纵轴方向）、单个气孔器面积、气孔器总面积较大,气孔密度及气孔指数较小。【结论与意义】假臭草叶片气孔特征表现可塑性,说明其对异质环境具有一定的生态适应能力。     

赤腹松鼠（Callosciurus erythralus）春季生境特征初步分析

2009年2月至5月,在广西龙江河畔对赤腹松鼠（C.erythralus）的春季生境特征进行了分析.野外共测量了57个10m×10m样方中的13个生态因子,并运用频次分析和主成分分析的方法,对赤腹松鼠的春季生境选择因子进行了分析.结果表明,赤腹松鼠春季生境的主要特征为：郁闭度良好,水源距离＜30m,坡度20～40°,避风性良好,坡向以东坡和南坡为主,坡位中坡位或上坡位,食物因子良好,人为干扰距离＜10m,海拔50～100m,乔木密度＜50株,乔木距离低于4m,灌木密度低于200株,灌木距离＜2m.影响赤腹松鼠春季生境选择的主要因子为郁闭度、避风性、坡度、坡位和灌木距离;次要因子为海拔、人为干扰距离、乔木距离、水源距离、乔木密度、灌木密度、食物丰富度、坡向.     

林下与全光下地枫皮叶片形态和光合特性的比较

测量了林下与全光下地枫皮的叶片形态和光合-光响应曲线,探讨光强对地枫皮的形态和生理特性的影响。结果表明：林下与全光下地枫皮叶片净光合速率（Pn）、气孔导度（Gs）、蒸腾速率（Tr）和水分利用效率（WUE）对光强的响应趋势均基本一致,但全光下的Pn、Gs和Tr值较高,林下WUE值较高。全光下地枫皮的最大净光合速率、光饱和点和光补偿点均极显著高于林下,但弱光下的量子效率无显著差异;林下地枫皮的叶长、叶宽、干物质重、叶面积和比叶面积等叶片形态参数均极显著大于全光。推断地枫皮为耐阴性较弱的阳生植物,其光合能力和光饱和点较低,是对干旱环境的适应性反应;全光下地枫皮叶片狭小降低了吸光面积,有利于避免过高光强对叶光合器官的损伤。     

Axial Water Flux Dynamics in Small Diameter Roots of a Fast Growing Tropical Tree

Field measurements of water flux in small diameter roots are important to the study of whole plant water transport systems. Miniature sap flow gauges were used to capture high resolution water flux patterns in small roots of 2 – 5 mm diameter and simultaneously in the canopy branches of a Eucalyptus saligna tree growing in Hawaii. The axial transport flux rates were then correlated with anatomical measurements to describe the internal hydraulic capacity of the tree. The daily patterns of water flux showed a strong coupling between the canopy and root systems and both systems were tightly synchronized with rapid fluctuations in photosynthetic photon flux density, vapour pressure deficit, and wind speed. When flow rates were normalized by the total vessel lumen area, branches had daily totals equivalent to the surface roots. Daily flows of water through surface roots were consistently 30% greater than through deep roots. Results of an experiment where a portion of the canopy was removed showed the decrease in water flux for all roots was in nearly direct proportion to the decrease in leaf area. The root anatomical measurements suggested a high capacity axial root water transport system with roots containing a smaller number of vessels per unit of sapwood area than branches but with vessel diameters twice that of the branches. However, relative conductivity values of roots and branches were similar and comparable to some of the highest values reported. Overall, the results suggested a highly efficient axial water transport system that would help to maintain a favorable plant water status for maximal stomatal opening. This revised version was published online in July 2006 with corrections to the Cover Date.     

Stomatal Density and Bio-water Saving

Bio-water saving is to increase water use efficiency of crops or crop yield per unit of water input. Plant water use efficiency is determined by photosynthesis and transpiration, for both of which stomata are crucial. Stomata are pores on leaf epidermis for both water and carbon dioxide fluxes that are controlled by two major factors： stomatal behavior and density. Stomatal behavior has been the focus of intensive research, while less attention has been paid to stomatal density. Recently, a number of genes controlling stomatal development have been identified. This review summarizes the recent progress on the genes regulating stomatal density, and discusses the role of stomatal density in plant water use efficiency and the possibility to increase plant water use efficiency, hence bio-water saving by genetically manipulating stomatal density.     

Epidermal conductance, stomatal density and stomatal size among genotypes of Sorghum bicolor (L.) Moench

Abstract. The ability of a plant to survive severe water deficits depends on its ability to restrict water loss through the leaf epidermis after stomata attain minimum aperture. At this stage, the rate of water loss is regulated by the epidermal conductance (g_c). Low g_c would be a useful selection criterion to identify genotypes with enhanced survival capability. Consequently, variation in g_c among Sorghum bicolor (L.) Moench genotypes was evaluated. Since there is little conclusive evidence linking g _c with leaf waxiness, alternative hypotheses relating g _c to stomatal trails were also examined. Epidermal conductance varied from 6.3 to 17.6mmol m⁻² s⁻¹ among sorghum genotypes. It was unrelated to stomatal pore length which varied with genotype and to pore depth which was similar for all genotypes measured. However, g _c, increased with increasing stomatal density. This indicates that stomatal density plays a direct role in water loss even at very low conductances. The association of low stomatal density with low g _c is consistent with the hypothesis that at the smallest stomata aperture, water loss from the epidermis above guard cell teichodes becomes a significant source of leaf water loss. Since low g _c is directly related to crop survival under severe water deficits, it is recommended that genotypes with low g _c. be selected using the selection criterion of stomatal density.     

A canopy-scale test of the optimal water-use hypothesis

Common empirical models of stomatal conductivity often incorporate a sensitivity of stomata to the rate of leaf photosynthesis. Such a sensitivity has been predicted on theoretical terms by Cowan and Farquhar, who postulated that stomata should adjust dynamically to maximize photosynthesis for a given water loss.
In this study, we implemented the Cowan and Farquhar hypothesis of optimal stomatal conductivity into a canopy gas exchange model, and predicted the diurnal and daily variability of transpiration for a savanna site in the wet–dry tropics of northern Australia. The predicted transpiration dynamics were then compared with observations at the site using the eddy covariance technique. The observations were also used to evaluate two alternative approaches: constant conductivity and a tuned empirical model.
The model based on the optimal water-use hypothesis performed better than the one based on constant stomatal conductivity, and at least as well as the tuned empirical model. This suggests that the optimal water-use hypothesis is useful for modelling canopy gas exchange, and that it can reduce the need for model parameterization.     

The control of stomata by water balance

It is clear that stomata play a critical role in regulating water loss from terrestrial vegetation. What is not clear is how this regulation is achieved. Stomata appear to respond to perturbations of many aspects of the soil-plant-atmosphere hydraulic continuum, but there is little agreement regarding the mechanism (or mechanisms) by which stomata sense such perturbations. This review discusses feedback and feedforward mechanisms by which hydraulic perturbations are putatively transduced into stomatal movements, in relation to generic empirical features of those responses. It is argued that a metabolically mediated feedback response of stomatal guard cells to the water status in their immediate vicinity ('hydro-active local feedback') remains the best explanation for many well-known features of hydraulically related stomatal behaviour, such as transient 'wrong-way' responses and the equivalence of hydraulic supply and demand as stomatal effectors. Furthermore, many curious phenomena that appear inconsistent with feedback, such as 'apparent feedforward' humidity responses and 'isohydric' behaviour (water potential homeostasis), are in fact expected to emerge from the juxtaposition of hydro-active local feedback and the well-known hysteretic and threshold-like effect of water potential on xylem hydraulic resistance.