干旱胁迫对4种植物蒸腾特性的影响

在温室条件下研究类芦等4种植物在干旱胁迫下的蒸腾特性。结果表明,在正常供水情况下,4种植物蒸腾速率和气孔阻力日变化规律明显;在干旱胁迫下,气孔阻力增大,蒸腾速率逐步降低,当水势下降到临界值时,气孔关闭,气孔阻力急剧增大,蒸腾速率降至最低。4种植物中,类芦的水分利用效率最高,其次为百喜草和香根草,五节芒最低。4种植物的抗旱力以类芦最强,百喜草和香根草其次,五节芒最弱。     

O_3和干旱胁迫对元宝枫叶片气孔特征的复合影响

利用开顶式气室(OTC)研究了O3胁迫(空气O3浓度加118μg·m-3)、干旱胁迫(土壤水分保持在田间持水量的40%~50%)以及O3与干旱复合胁迫(空气O3浓度加118μg·m-3和土壤水分保持在田间持水量的40%~50%)对元宝枫(Acer truncatum Bunge.)叶片气孔特征的影响。结果表明:O3胁迫41 d时气孔密度变化不明显,62和110 d时显著增加(P0.05);干旱胁迫和复合胁迫下气孔密度增加显著(P0.05);O3、干旱及复合胁迫处理110 d气孔面积、周长、长度和宽度明显减小(P0.05),气孔指数明显增加(P0.05);各胁迫气孔开度均明显减小(P0.05),O3和干旱对气孔开度影响存在协同作用,干旱减小了O3胁迫下的气孔开度。O3和干旱胁迫下气孔特征响应不同表明影响气孔的机制不同,O3对气孔影响具有累积效应,干旱可限制O3摄入量从而减小了O3对植物的伤害。     

甘草叶片形态结构和光合作用对干旱胁迫的响应

叶片结构在植物防御生物和非生物胁迫方面起着重要的作用,可通过合成、储存和分泌次生代谢产物提高植物抗性。以甘草幼苗为试材,采用盆栽控水自然干旱法,探讨叶片光合作用、气孔微形态和腺体形态对干旱胁迫的响应。结果表明:①随着干旱胁迫程度的加剧,叶片净光合速率(Pn)、气孔导度(Gs)和蒸腾速率(Tr)均呈先升高后降低的趋势;其中胞间CO2浓度(Ci)在重度干旱胁迫(severe stress,SS)时迅速增高。②随着干旱胁迫程度的加剧,叶片总气孔密度和气孔开张比呈先增大后减小的趋势;而气孔开张宽度呈逐渐减小的趋势。③随着干旱胁迫程度的加剧,叶片上表皮和下表皮腺体密度总数整体上呈增大的趋势,腺体颜色随着干旱胁迫程度的加剧逐渐加深,形状出现不规则褶皱和内陷。总之,甘草叶片表面的腺体特征参与抗旱逆境调节,从而避免干旱胁迫对甘草植株的伤害;在SS下,胁迫程度加速了气孔细胞的程序性死亡(PCD),甘草幼苗失去抗旱能力。     

臭氧与干旱对植物复合影响的研究进展

地表臭氧(O_3)浓度和干旱频率的持续增加成为限制植物生长的重要因素。O_3通过气孔扩散进入植物组织内部,产生并积累活性氧(ROS)自由基,促发细胞程序性死亡。干旱破坏植物抗氧化系统对ROS的解毒和修复功能,导致ROS累积。两种胁迫对植物的影响都是积累ROS并引发氧化胁迫,使植物的光合作用和生理代谢机能受到限制,最终阻碍植物生长,导致生物量降低。然而,O_3和干旱胁迫对植物的复合效应可能是协同加重植物损伤,也可能是拮抗减轻植物伤害,二者的交互影响存在复杂的作用过程。一方面,O_3引起气孔响应滞后甚至失灵,使植物对于两种胁迫的响应变得迟钝,进而加重植物的蒸散失水和O_3毒害。另一方面,干旱使植物气孔关闭,从而降低对O_3的吸收量和水分蒸发,但长期干旱限制CO2的吸收,最终导致植物的生长受限。植物的响应过程不仅取决于两种胁迫作用的先后次序和持续时间,而且受到植物本身生理代谢差异的影响。该文结合国内外研究,从气孔、光合碳代谢、抗氧化系统和生长发育等方面阐述了O_3和干旱胁迫对植物代谢调节和生长发育的复合影响,并提出了未来研究的发展方向。     

液压信号与植物水分胁迫信号传递研究（综述）

植物根部受到干旱胁迫时,由茎部产生的液压信号使茎部作出反应.液压传递根到茎之间的水分胁迫信号.土壤干旱引起茎中场所产生液压反应,然后产生ABA信号,引起气孔关闭.在不同植物中,减弱液压反应,阻止干旱信号的长距离传导,气孔不能关闭.     

前期水淹对牛鞭草后期干旱胁迫光合生理响应的影响

水淹和干旱是限制植物生长的两种主要环境因子。三峡库区消落带由于其特殊的地形条件和人工水文节律,呈现以年度为周期的“水淹-落干”交替变化的水文变动特征,在消落带生长的植物因此受到水淹和干旱交替胁迫的双重影响。为了探究库区蓄水对消落带植被干旱耐受性的影响,以当年生牛鞭草扦插苗为试验对象,设置对照组(CK)、表土水淹组(SF)、全淹组(TF)、对照-干旱组(CD)、表土水淹-干旱组(SFD)、全淹-干旱组(TFD)6个处理组,研究不同水分处理对牛鞭草光合特性的影响。结果表明:(1)水淹和干旱胁迫均对牛鞭草光合特性造成显著影响;(2)水淹胁迫阶段,与CK组相比,牛鞭草SF和TF组净光合速率、气孔限制值和水分利用效率显著下降,胞间CO_2浓度显著上升;(3)干旱胁迫阶段,牛鞭草CD和SFD组净光合速率、气孔导度、胞间CO_2浓度和蒸腾速率等光合参数显著低于CK组,TFD组净光合速率、气孔导度、胞间CO_2浓度和蒸腾速率等指标与CK组无显著差异;(4)复水阶段,各处理组净光合速率、气孔导度、胞间CO_2浓度和蒸腾速率等指标均与CK组无显著差异。研究表明,前期水淹并未增加牛鞭草对后期干旱胁迫的敏感性,牛鞭草对水淹和干旱胁迫均具有较好的耐受性,有助于牛鞭草对库区消落带生境变化的适应性。     

干旱胁迫对香紫苏生理特性和光合特性的影响研究

目的:研究土壤干旱胁迫对香紫苏生理特性和光合特性的影响.方法:以香紫苏(Salvia sclarea L)的功能叶为研究对象,分实验组和对照组进行盆栽实验,在随后的5天中,分别测定两组香紫苏叶片中脯氨酸、丙二醛和叶绿素的含量,同时测定并分析蒸腾速率、气孔导度、净光合速率、细胞间隙CO2浓度的变化情况.结果:随着干旱胁迫程度的加重,香紫苏叶片内脯氨酸的含量总体趋势增加,与干旱胁迫的时间正相关;而叶片中丙二醛的含量变化不明显;干旱胁迫时,叶片内叶绿素的含量升高,在第四天时达到最大值,随干旱胁迫的加重,叶绿素含量又呈下降的趋势.其蒸腾速率变化曲线呈双峰型,随干旱时间的延长,蒸腾速率明显下降;香紫苏气孔导度变化曲线为单峰型,当干旱胁迫加重时,香紫苏气孔导度下降明显,气孔导度与胞间CO2浓度呈负相关,净光合速率变化与气孔导度变化曲线较为一致.结论:香紫苏生理因子和光合作用对干旱胁迫有一定的适应能力,但重度胁迫会对其造成严重影响.     

植物应对干旱胁迫的气孔调节

气孔是植物控制叶片与大气之间碳、水交换的重要门户,植物的生长和生存都依赖于叶片气孔对碳获取和水散失的调控.因此,气孔调节机理研究与气孔导度模型研发是精确模拟陆地生态系统碳、水循环过程不可或缺的内容.近年来,随着气候变化的加剧,干旱事件愈发频繁,对植物的存活、生长和分布产生深刻影响.为了深入理解植物碳-水耦合机理过程、预测全球变化下植物及群落的动态,开展植物应对干旱胁迫的气孔调节研究尤为重要.本文综述了植物在干旱胁迫条件下气孔调节机制和模型研究进展.首先阐述了植物气孔对干旱胁迫的主动调节与被动调节,讨论了气孔调节的演化过程,包括蕨类和石松类植物的被动水力调节、被子植物的主动调节和裸子植物的双重调节机制,认为裸子植物的气孔调节方式是植物进化过程中介于蕨类、石松类植物和被子植物之间的一种重要过渡类型.然后分析了气孔调节与水力调节的关系,讨论了“植物水势和气孔导度解耦”问题中存在的争议.之后介绍了基于水分利用效率假说和最大碳增益假说所建立的气孔导度优化模型的应用,并指出后者有更强的预测能力和应用前景.最后,为了有效减少植被对气候变化响应预测中的不确定性,提出了2个亟待开展的研究问题:将植物叶片的气孔调节功能研究由个体扩展到生态系统甚至更大尺度,改进陆地生态系统碳水循环机理模型;量化气孔调节的主动水力反馈过程,修正植物气孔功能水力模型.     

干旱胁迫下水杨酸对银沙槐子叶表皮气孔开度的影响

以沙生植物银沙槐似mmodendronargenteum）幼苗为实验材料,采用PEG处理和PEG＋SA综合处理两种方式,其中SA包括不同浓度、不同时间和不NpH值3种处理,显微观察干旱胁迫处理、干旱胁迫下外源水杨酸处理对子叶表皮气孔开度的影响,以探讨干旱与气孔运动的关系。结果表明：干旱胁迫处理后,气孔运动的趋势是随着PEG浓度的增加,孔径逐渐减小。在干旱胁迫下,sA不同浓度处理后,气孔孔径在相同PEG浓度下随着sA浓度的增加而减小;随sA处理时间的不同,气孔开度随着时间的延长而变小,且在不同PEG浓度（0、10％和20％）条件下,气孔开度的变化与sA处理时间均呈显著（R=-0．998＊＊、R=-0．955＊和R=-0．983＊）负相关;sA溶液pH值越低,对气孔开度的抑制作用越明显。可见,干旱胁迫后能减小银沙槐气孔孔径的大小,在干旱胁迫处理下施加SA处理,sA对银沙槐气孔开度的抑制作用加剧,说明sA在提高银沙槐的抗旱性方面起到了一定作用。     

脱落酸调节植物抵御水分胁迫的机制研究

干旱、盐渍、低温等均可导致植物可利用水分的亏缺,表现为水分胁迫。植物感受到水分胁迫,诱导脱落酸（abscisic acid,ABA）生物合成。ABA可通过促使气孔关闭或抑制气孔开放,使作物尽可能地降低蒸腾失水,以抵御水分胁迫。该文就植物激素ABA及其下游信号过氧化氢（hydrogenperoxide,H2O2）、一氧化氮（nitric oxide,NO）以及Ca2＋等在植物气孔运动调节方面的研究进展进行概述,以构建水分胁迫下ABA调节植物气孔运动的可能模式。     

干旱下植物气孔运动的调控

概述了植物气孔对大气干旱和土壤干旱的反应,认为植物气孔对大气干旱的反应并不是一种反馈机制;并就干旱条件下植物气孔运动的水力学和化学信号调控机制进行了简要论述,认为虽然化学信号调控干旱下气孔运动更为广泛,但ABA不是唯一的化学信号,水分关系影响了信号的产生、运转和气孔对信号的敏感性,干旱条件下水力学和化学信号共同调控着植物的气孔运动。     

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.     

丛枝菌根真菌对杨树生长、气孔和木质部微观结构的影响

植物气孔与木质部导管及纤维的功能直接关系着植物的水分利用, 进而影响植物的生长。为研究丛枝菌根真菌(AMF)对杨树抗旱性的影响, 采用温室盆栽的方法, 研究两种水分条件下, 接种根内球囊霉(Rhizophagus irregularis)对速生杨107 Populus × canadensis (P. nigra × P. deltoides) ‘Neva’气孔及木质部微观结构的影响。结果表明: AMF的侵染显著提高了杨树幼苗地上和地下部分生物量, 对叶片气孔长度、茎部导管细胞直径和纤维细胞长度也有促进作用。AMF对生物量和导管细胞直径的增加幅度表现出干旱条件下>正常水分条件下, 而对气孔长度的提高幅度表现出干旱条件下<正常水分条件下。正常水分条件下, AMF增加了杨树叶片的气孔密度, 减小了纤维细胞直径, 对相对水分饱和亏缺无影响; 干旱条件下, AMF增加了纤维细胞直径, 降低了相对水分饱和亏缺, 对气孔密度无影响。综上所述, 干旱条件下, AMF对导管水分传输能力的促进作用明显增加, 而对气孔蒸腾能力的促进作用有所减少, 从而更利于杨树在遭遇干旱时保持水分, 减少干旱对菌根杨树造成的水分亏缺, 提高菌根杨树对干旱的耐受性。     

Plant resistance to drought depends on timely stomatal closure

Stomata play a significant role in the Earth's water and carbon cycles, by regulating gaseous exchanges between the plant and the atmosphere. Under drought conditions, stomatal control of transpiration has long been thought to be closely coordinated with the decrease in hydraulic capacity (hydraulic failure due to xylem embolism). We tested this hypothesis by coupling a meta‐analysis of functional traits related to the stomatal response to drought and embolism resistance with simulations from a soil–plant hydraulic model. We report here a previously unreported phenomenon: the existence of an absolute limit by which stomata closure must occur to avoid rapid death in drought conditions. The water potential causing stomatal closure and the xylem pressure at the onset of embolism formation were equal for only a small number of species, and the difference between these two traits (i.e. safety margins) increased continuously with increasing embolism resistance. Our findings demonstrate the need to revise current views about the functional coordination between stomata and hydraulic traits and provide a mechanistic framework for modeling plant mortality under drought conditions.     

The plant-specific actin binding protein SCAB1 stabilizes actin filaments and regulates stomatal movement in Arabidopsis

Microfilament dynamics play a critical role in regulating stomatal movement; however, the molecular mechanism underlying this process is not well understood. We report here the identification and characterization of STOMATAL CLOSURE-RELATED ACTIN BINDING PROTEIN1 (SCAB1), an Arabidopsis thaliana actin binding protein. Plants lacking SCAB1 were hypersensitive to drought stress and exhibited reduced abscisic acid-, H(2)O(2)-, and CaCl(2)-regulated stomatal movement. In vitro and in vivo analyses revealed that SCAB1 binds, stabilizes, and bundles actin filaments. SCAB1 shares sequence similarity only with plant proteins and contains a previously undiscovered actin binding domain. During stomatal closure, actin filaments switched from a radial orientation in open stomata to a longitudinal orientation in closed stomata. This switch took longer in scab1 plants than in wild-type plants and was correlated with the delay in stomatal closure seen in scab1 mutants in response to drought stress. Our results suggest that SCAB1 is required for the precise regulation of actin filament reorganization during stomatal closure.     

外源脯氨酸对自然干旱下白刺叶片气孔的影响

以大田环境内多年生荒漠植物白刺（Nitraria tangutorum）为研究对象,采用扫描电子显微镜对不同外源脯氨酸质量浓度处理下白刺叶片气孔进行微观结构观察并测定叶片表皮气孔器长度、宽度、面积和密度,比较同一指标在不同脯氨酸质量浓度处理下的差异。采用室外试验研究自然干旱胁迫下喷施质量浓度为50、100、150、200、250 mg·L^-1的外源脯氨酸（Proline,缩写为Pro或P）研究白刺的耐旱性。观察结果显示,白刺叶片气孔保卫细胞为肾形,气孔在叶片表皮随机分布,气孔器多为无规则型,气孔呈椭圆形且叶片表面蜡质较少,并且叶片细胞出现褶皱和下陷。与对照相比,在不同质量浓度处理和不同采样时间时,气孔长度、宽度及面积下降,气孔密度增大,随着浓度的升高以及采样时间的变化,气孔长度整体呈现下降趋势,当浓度达到200 mg·L^-1时,下降幅度增大,宽度和面积整体呈现先降低后升高再降低的趋势,而气孔密度则整体呈现上升趋势,其中,气孔长度、宽度和面积在质量浓度为50 mg·L^-1、采样第1天时达到最大值,气孔密度在质量浓度为150 mg·L^-1、采样第9天时达到最大值。此外,气孔长度在不同浓度脯氨酸处理下第1、6和9天存在显著差异,而气孔宽度和面积差异不显著,除第0天外,气孔密度因脯氨酸质量浓度不同均差异显著（P<0.05）。研究结果表明,自然干旱胁迫下对白刺叶片喷施不同质量浓度的外源脯氨酸,可减小气孔长度、宽度和面积,而增加气孔密度,以期为白刺抗旱提供理论依据。     

植物气孔气态失水与SPAC系统液态供水的相互调节作用研究进展

讨论了植物气孔气态失水与SPAC系统液态供水相互作用研究领域的一些重要现象和行为.当植物水力信号和化学信号共同作用促进气孔对叶水势的调节时,植物对叶水势的调节表现为等水行为.气孔对环境湿度变化响应的反馈机制可用来解释土壤干旱条件下气孔和光合的午休现象,以及气孔导度和水流导度之间的相关关系;而气孔对环境湿度变化响应的前馈机制,则可用来解释气孔导度对大气 叶片间水汽饱和差的滞后反应.植物最大限度地利用木质部传输水分的策略,要求气孔快速响应以避免木质部过度气穴化和短时间内将气穴逆转的相应机制.     

Interactive effects of ozone and drought stress on plants: A review

Ground-level ozone (O₃) and drought are two key factors limiting plant growth. O₃ can enter into the plant tissue through the stomata, then causing the formation of reactive oxygen species (ROS) which inspires programmed cell death. Drought usually induces the accumulation of ROS due to damage to antioxidant systems of plants. The effects of two kinds of stress on plants are similar due to the accumulation of ROS, resulting in reduced photosynthesis rate and physiological metabolism, eventually decreased plant growth and biomass. Nevertheless, O₃ and drought interacts synergistically to accumulate detrimental effects or antagonistically to reduce harmful effects. Actually, it is complex interactive process between O₃ and drought. On the one hand, O₃ triggers stomatal sluggishness or even dysfunction, which exacerbates water transpiration of leaves, water loss from plants and further O₃ phytotoxicity. On the other hand, drought induces stomatal closure, and thus protecting plants against the O₃ influx and evaporation of water. However, prolonged drought could limit the uptake of CO₂ and thus result in reduced plant growth. The response of plants to both O₃ and drought not only depends on the occurring sequence and duration of any factor but also rely on the difference in physiological metabolism of the plant itself. The interactive effects of O₃ and drought on stomatal characteristics, photosynthetic carbon mechanism, antioxidant response and growth development are reviewed in this paper and the aspects to be further studied are also suggested.     

Farnesol-like antitranspirant activity and stomatal behaviour in maize and Sorghum lines of differing drought tolerance

Abstract Potted seedlings of four lines of maize and Sorghum of differing drought tolerance were subjected to a single soil drying cycle and were only rewatered when the plants showed the first signs of wilting. Other plants remained well-watered throughout the experimental period. As plant water potentials decreased in the unwatered plants of three of the lines investigated (Sorghum Piper and M35-1, V-4146 and maize Farz 27), endogenous levels of farnesol-like antitranspirants increased. Closure of stomata correlated well with the increase in endogenous antitranspirant. In the fourth line (Sorghum M35-1, V-4184), stomata did not close as the level of plant water stress increased, although leaf diffusion resistance of even the well-watered plants of this line was quite high. In this line, there was no consistent relationship between plant water stress and antitranspirant level or between stomatal behaviour and antitranspirant level. The involvement of farnesol-like antitranspirants in the control of stomatal behaviour in water-stressed plants is discussed.     

The effect of air pollution on the mechanism of stomatal control

Summary A long-term field experiment permanently measuring gas exchange in the top of a 70-year-old spruce, continued for through the 1990 growing season. Two gas exchange chambers were run simultaneously under identical climatic conditions. One of two similar twigs was exposed to ambient air whereas the other received pure air. These experiments aimed to examine the ability of the stomata to control water balance, comparing pure and ambient air. This was done not only in natural climatic conditions but also in experimental, specifically maintained stress situations. Special care was taken to ensure that only steady state values of stomatal responses are related to the environmental stimuli. During a drought period lasting several weeks, overshooting transpiration values were documented for the ambient air. The two twigs do not merely differ in their control capacity, but the behaviour of the stomata in ambient air deviates from the norm. The increasingly uncontrolled water losses during the drought period have a negative effect on photosynthetic capacity. The influence of water deficit on stomatal response to other environmental factors (light, CO₂) is shown. Due to deficient control quality of the stomata lower stress tolerance in the face of drought is suggested in ambient air as compared with pure air. By tracing dysfunctions to structural changes in the cell walls of the stomatal apparatus, a mechanism is described explaining forest decline under the combined influence of air pollutants and drought stress.