Modulation of Root Signals in Relation to Stomatal Sensitivity to Root-sourced Abscisic Acid in Drought-affected Plants

Stomatal sensitivity to root signals induced by soil drying may vary between environments and plant species. This is likely central role in root to shoot signaling. pH and hydraulic signals may interact with ABA signals and thus, jointly regulate stomatal responses to changed soil water status. pH itself can be modified by several factors, among which the chemical compositions In the xylem stream and the live cells surrounding the vessels play crucial roles. In addition to the xylem pH,more attention should be paid to the direct modulation of leaf apoplastic pH, because many chemical compositions might strongly modify the leaf apoplastlc pH while having no significant effect on the xylem pH. The direct modulation of the ABA signal intensity may be more important for the regulation of stomatal responses to soil drying than the ABA signal per se.The ABA signal is also regulated by the ABA catabolism and the supply of precursors to the roots If a sustained root to shoot communication of soil drying operates at the whole plant level. More importantly, ABA catabolism could play crucial roles In the determination of the fate of the ABA signal and thereby control the stomatal behavior of the root-sourced ABA signal.     

干旱胁迫下氮素营养与根信号在气孔运动调控中的协同作用

干旱胁迫下根系与地上部分之间的信息传递可使植物叶片及时感知土壤水势变化,从而使植物在没有真正受到干旱伤害时即可做出主动、快速的抗旱应答反应,而在这一过程中,脱落酸（abscisic acid,ABA）和pH起着关键的作用。本研究表明。干旱胁迫下鸭趾草（Commelina communis L．）、番茄（Lycopersicon esculentum Mill．）和向日葵（Helianthus annuus L．）木质部汁液中pH的变化很不相同,且该pH变化和木质部汁液中硝态氮离子浓度的变化没有直接的关系;然而,饲喂实验表明,无论对于何种植物,蒸腾流中硝态氮离子浓度的增加都可有效地增加气孔对ABA的敏感度;分根实验进一步表明,土壤中硝态氮营养的增加可明显提高气孔对根信号的敏感度。以上结果说明,氮素营养可以和根信号相互作用共同操纵气孔运动。     

Deficient Glutathione in Guard Cells Facilitates Abscisic Acid-Induced Stomatal Closure but Does Not Affect Light-Induced Stomatal Opening

We investigated the role of glutathione (GSH) in stomatal movements using a GSH deficient mutant, chlorinal-1 (ch1-1). Guard cells of ch1-1 mutants accumulated less GSH than wild types did. Light induced stomatal opening in ch1-1 and wild-type plants. Abscisic acid (ABA) induced stomatal closure in ch1-1 mutants more than wild types without enhanced reactive oxygen species (ROS) production. Therefore, GSH functioned downstream of ROS production in the ABA signaling cascade.     

植物ABA-H2O2介导的气孔关闭

本文综述了脱落酸作为根源信号物质经由木质部被传递到叶片,经重新分配再与脱落酸受体结合,然后刺激气孔开放因子,调节烟酰胺腺嘌呤二核苷酸磷酸氧化酶等关键酶活性产生过氧化氢,过氧化氢可使胞质碱化并刺激钙离子通道使钙离子内流,活化阴离子通道使阴离子外流,最终导致气孔关闭的一系列过程。该过程涉及到的因子包括：脱落酸受体、气孔开放因子、磷脂酰环己六醇、分裂原激活蛋白激酶、烟酰胺腺嘌呤二核苷酸磷酸氧化酶、Ca^（2＋）、pH、一氧化氮等。     

植物磷脂酸的特性及其在ABA诱导气孔运动中的作用

磷脂酸(PA)是应答多种生理过程的第二信使, 其作为一个脂质信号快速积累从而响应多种环境。PA主要通过磷脂酶D (PLD)和磷脂酶C/甘油二酯激酶(PLC/DGK)途径产生。基于PLDs的生化特性、激活机制以及在不同类型胁迫下被激活的特定同种型的差异, 不同类型胁迫下会产生特定分子种组成的PA。PA在多种环境下起信号转导作用, 在调节气孔运动中, PA的作用模式主要是通过与多种蛋白结合, 激活或抑制这些蛋白的活性, 进而执行其信使功能。该文主要综述PA的生化特性以及信号途径中PA互作蛋白的研究进展, 并提出PA研究中亟待解决的问题及今后的重点研究方向。     

Modeling the Calcium Signaling in Stomatal Guard Cells under the Action of Abscisic Acid

A theoretical model of calcium signaling is presented that simulates oscillations of cytoplasmic calcium concentration ([Ca²⁺]_cyt) in stomatal guard cells under the action of abscisic acid. The model is based on the kinetics of inositol 1,4,5-trisphosphate-sensitive calcium channels of endoplasmic reticulum and cyclic ADP-ribose-sensitive calcium channels of the tonoplast. The operation of two energy-dependent pumps—the Ca²⁺-ATPase of the endoplasmic reticulum and the Ca²⁺/H⁺ antiporter of the tonoplast—is also included in the model. It is shown that the removal of excessive Ca²⁺ from the cytoplasm by the tonoplast Ca²⁺/H⁺ antiporter is the main factor accounting for generation of [Ca²⁺]_cyt oscillations at a wide range of ABA concentrations (0.01–1 M). The long period of [Ca²⁺]_cyt oscillations in plant cells is explained by a slow release from inhibition of inositol 1,4,5-trisphosphate-gated calcium channels.     

保卫细胞的ABA信号转导

植物激素脱落酸(ABA)调节植物体多种生理过程,尤其在一些逆境条件下,植物体中ABA大量合成,诱导气孔关闭,从而有效地调控植物体内的水分平衡.尽管人们对ABA诱导气孔关闭作用已得到共识,但有关信号转导的细节还很不清楚.该文简要介绍了研究气孔保卫细胞信号转导途径的相关技术以及与ABA信号转导直接相关的ABA受体、第二信使、蛋白质磷酸化和离子通道调节等方面的最新妍究进展.并在前人研究工作的基础上,勾画出气孔保卫细胞ABA、H2O2的信号转导模式图.     

Abscisic acid concentrations and fluxes in droughted conifer saplings

We present the first study of abscisic acid (ABA) concentrations and fluxes in the xylem sap of conifers during a drought cycle. In both Pinus sylvestris and Picea sitchensis the concentration of ABA in the sap rose 11-fold as the drought progressed. There were clear diurnal trends in this concentration, which reached its maximum (6–8.ininol ABA m^?3) near the middle of the day. The fluxes of ABA were calculated by multiplying the xylem ABA concentration by the sap flow rate. The ABA fluxes in the droughted plants in the middle of the day were usually no higher than those of the controls, as a result of the very low sap flow in the droughted plants at that time. However, the ABA flux in the droughted plants was higher than in the controls in the morning, and we postulate that the stomata are responding to these ‘morning doses’ Stomatal conductance, g_s, could not be related statistically to leaf turgor or to the ABA flux. However, £s did display a negative exponential relationship with ABA concentration in the xylem. Pinus displayed more acclimation to drought than Picea, Its ABA concentration rose and its stomatal conductance fell at day 6 of the drought, as opposed to day 17 for Picea, and its osmotic potential fell during the drought treatment.     

Water Relations of Cotton Plants under Nitrogen Deficiency: V. Environmental Control of Abscisic Acid Accumulation and Stomatal Sensitivity to Abscisic Acid

Suboptimal N nutrition increased the water potential for stomatal closure in water stressed cotton (Gossypium hirsutum L.) leaves. This increased sensitivity to water stress had two components, increased accumulation of abscisic acid (ABA) and increased apparent stomatal sensitivity to ABA. Low N increased the threshold water potentials for stomatal closure and ABA accumulation by about 4 bars and 2 bars, respectively. Low N also greatly increased stomatal response to low concentrations of exogenous ABA applied to excised leaves through the transpiration stream. In low N leaves, kinetin decreased stomatal response to ABA to the level observed with high N leaves. Kinetin by itself had little effect on stomata, nor did it alter stomatal response to ABA in high N leaves. The results suggest a cytokinin-ABA balance which is altered by suboptimal N nutrition to favor stomatal closure during stress.

Ambient temperature and N nutrition interacted to alter stomatal response to water stress. Stress-induced ABA accumulation and apparent stomatal sensitivity to ABA were independently affected. The effects of each treatment, and their interaction, could be explained as the net result of changes in both accumulation and apparent sensitivity. Although the results document environmental control of stomatal response to ABA, either altered partitioning of ABA between active and inactive pools, or altered sensitivity of the guard cells, could account for the data.

     

Developmental Priming of Stomatal Sensitivity to Abscisic Acid by Leaf Microclimate

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Stomatal Reactions of Two Different Maize Lines to Osmotically Induced Drought Stress

Two maize lines differing in drought resistance were grown at different drought stress induced by polyethylene glycol (PEG 10 000) solutions with osmotic potentials of –0.20, –0.40 and –0.80 MPa in the semipermeable membrane system. During the five days soil water content decreased (from 0.43 to 0.29, 0.25 and 0.23 g cm^–3 for three PEG solutions, respectively) as well as leaf water potentials (_w; from – 0.54 to –0.76, –1.06 and –1.46 MPa). These values were not significantly different between the investigated lines, indicating that a controlled and consistent soil moisture stress was achieved. Soil drying induced an increase in the ABA content of leaves and xylem of both lines and the effects on stomatal conductance were greater in drought susceptible line (B-432) compared to drought resistant line (ZPBL-1304). To test possible difference in stomatal sensitivity to xylem ABA between lines and to assess any ABA vs. _w interaction, roots were fed with 10, 50 and 100 mmol m^–3 ABA solutions in another set of experiments. These results showed that manipulation of xylem ABA affected stomata of both lines similarly. Comparison of stomatal sensitivity to drought-induced and applied ABA demonstrated that drought treatment affected stomata of investigated lines by differentially increasing their sensitivity to xylem ABA, thus confirming an interaction between chemical signalling and hydraulic signalling.     

Stomatal movement in response to long distance-communicated signals initiated by heat shock in partial roots of Commelina communis L.

The systematic or long-distance signal transmission plays crucial roles in animal lives. Compared with animals, however, much less is known about the roles of long-distance signal communication in plant lives. Using the model plant Commelina communis L., we have probed the root to shoot communication mediated by heat-shock signals. The results showed that a heat shock of 5 min at 40℃ in partial roots, i.e. half or even 1/4 root system, could lead to a significant decrease in stomatal conductance. The regulation capability depends on both heat shock temperature and the amount of root system, i.e. with higher temperature and more roots stressed, the leaf conductance would decrease more significantly. Interestingly, the stomatal regulation by heat shock signal is in a manner of oscillation: when stomata conductance decreased to the lowest level within about 30 min, it would increase rapidly and sometimes even exceed the initial level, and after several cycles the stomata conductance would be finally stabilized at a lower level. Feeding xylem sap collected from heat-shocked plants could lead to a decrease in stomata conductance, suggesting that the heat shock-initiated signal is basically a positive signal. Further studies showed that heat shock was not able to affect ABA content in xylem sap, and also, not able to lead to a decrease in leaf water status, which suggested that the stomatal regulation was neither mediated by ABA nor by a hydraulic signal. Heat shock could lead to an increase in xylem sap H₂O₂ content, and moreover, the removal of H₂O₂ by catalase could partially recover the stomatal inhibition by xylem sap collected from heat-shocked plants, suggesting that H₂O₂ might be able to act as one of the root signals to control the stomatal movement. Due to the fact that heat-shock and drought are usually two concomitant stresses, the stomatal regulation by heat-shock signal should be of significance for plant response to stresses. The observation for the stomatal regulation in an oscillation manner by presently identified new signals should contribute to further understanding of the mystery for the pant systematic signaling in response to stresses.     

脱落酸抑制蓝光依赖气孔开放运动的生理机制分析

以蚕豆(Vicia fabaL.)为材料,采用表皮条生物学分析技术、远红外成像技术以及电生理膜片钳技术,研究逆境信号脱落酸(ABA)和蓝光信号对蚕豆叶片表皮气孔运动及质膜K+通道的影响,以探讨ABA调节蓝光诱导的气孔开放运动的生理机制.结果表明:(1)100μmol?m-2?s-1蓝光能显著诱导气孔开放,该效应可被ABA以浓度依赖的方式抑制,并以10μmol?L-1ABA抑制效应最为明显.(2)100μmol?m-2?s-1蓝光处理能明显促使蚕豆叶面温度下降,而10μmol?L-1ABA可显著阻止蓝光诱导的蚕豆叶面温度下降.(3)100μmol?m-2?s-1蓝光可明显激活保卫细胞质膜内向K+通道,处理5 min后内向K+电流增加58%;对于1和10μmol?L-1ABA预处理蚕豆保卫细胞原生质体,在蓝光处理5 min后其内向K+电流增加值分别被抑制25%和51%,但10μmol?L-1ABA并不明显抑制壳梭孢菌素(质膜H+-ATP酶永久性激活剂)诱导的保卫细胞内向K+电流上升.研究发现,逆境信号ABA可能主要通过抑制蓝光信号转导中质膜H+-ATP酶上游位点,阻断蓝光激活的保卫细胞质膜内向K+通道,抑制蓝光诱...     

Immunogold Localization and Quantification of Cellular and Subcellular Abscisic Acid, Prior to and During Drought Stress

An Immunogold labeling procedure and experimental data are presented, which demonstrate that antibodies produced against a bovine serum albumin-abscisic acid conjugate can be used both to characterize the cellular and subcellular localization of abscislc acid (ABA), and to permit quantitative comparisons of this hormone in the subcellular compartments prior to and at times of drought stress. At the control leaf water potential (approximately -0.45 MPa), a quantitatively similar positive labeling pattern was observed in the chloroplasts and apoplast. A twofold drought stress-induced increase in the apoplastic ABA concentration was observed in the drought stressed leaf tissue (i.e., at a leaf water potential of approximately -1.55 MPa), while the ABA concentration in the chloroplasts did not differ from that of the controls. Three histochemical controls and the physiological observations validated the specificity of the procedure. Based on the labeling patterns we observed and literature cited, the validity of the hypothesis that drought stress induces a release of chloroplastic ABA is questioned. We interpreted our results as providing indirect evidence for a drought stress-induced root source origin for the increased apoplastic ABA concentrations.     

Stomatal movement in response to long distance-communicated signals initiated by heat shock in partial roots of Commelina communis L.

The systematic or long-distance signal transmission plays crucial roles in animal lives. Compared with animals, however, much less is known about the roles of long-distance signal communication in plant lives. Using the model plant Commelina communis L., we have probed the root to shoot communication mediated by heat-shock signals. The results showed that a heat shock of 5 min at 40°C in partial roots, i.e. half or even 1/4 root system, could lead to a significant decrease in stomatal conductance. The regulation capability depends on both heat shock temperature and the amount of root system, i.e. with higher temperature and more roots stressed, the leaf conductance would decrease more significantly. Interestingly, the stomatal regulation by heat shock signal is in a manner of oscillation: when stomata conductance decreased to the lowest level within about 30 min, it would increase rapidly and sometimes even exceed the initial level, and after several cycles the stomata conductance would be finally stabilized at a lower level. Feeding xylem sap collected from heat-shocked plants could lead to a decrease in stomata conductance, suggesting that the heat shock-initiated signal is basically a positive signal. Further studies showed that heat shock was not able to affect ABA content in xylem sap, and also, not able to lead to a decrease in leaf water status, which suggested that the stomatal regulation was neither mediated by ABA nor by a hydraulic signal. Heat shock could lead to an increase in xylem sap H₂O₂ content, and moreover, the removal of H₂O₂ by catalase could partially recover the stomatal inhibition by xylem sap collected from heat-shocked plants, suggesting that H₂O₂ might be able to act as one of the root signals to control the stomatal movement. Due to the fact that heat-shock and drought are usually two concomitant stresses, the stomatal regulation by heat-shock signal should be of significance for plant response to stresses. The observation for the stomatal regulation in an oscillation manner by presently identified new signals should contribute to further understanding of the mystery for the pant systematic signaling in response to stresses.     

Stomatal movement in response to long distance-communicated signals initiated by heat shock in partial roots of Commelina communis L.

It is well known that animals are able to positively respond to environmental stimuli, thereby avoiding or reducing the possible injures or impacts on them, and the base for the rapid and positive responses is long- distance signal transmission mediated b…     

Stomatal response to drying soil in relation to changes in the xylem sap composition of Helianthus annuus. II. Stomatal sensitivity to abscisic acid imported from the xylem sap

Sunflower plants [Helianthus annuus L.) were subjected to soil drought. Leaf conductance declined with soil water content even when the shoot was kept turgid throughout the drying period. The concentration of abscisic acid in the xylem sap increased with decreasing soil water content. No general relation could be established between abscisic acid concentration in the xylem sap and leaf conductance due to marked differences in the sensitivity of leaf conductance of individual plants to abscisic acid from the xylem sap. The combination of these results with data from Gollan, Schurr & Schulze (1992, see pp. 551–559, this issue) reveals close connection of the effectiveness of abscisic acid as a root to shoot signal to the nutritional status of the plant.     

脱落酸与植物细胞的抗氧化防护

水分胁迫是一种影响植物生长发育、限制植物产量的重要胁迫因子.植物能够通过感知刺激、产生和传导信号、启动各种防护机制来响应与适应水分胁迫.植物激素脱落酸(ABA)作为一种胁迫信号,在调节植物对水分胁迫的反应中起着重要的作用.ABA不仅能诱导气孔关闭,而且能诱导编码耐脱水蛋白的基因表达.正在增加的证据显示,ABA增强水分胁迫的耐性与其诱导抗氧化防护系统有关.本文综述了ABA在诱导活性氧(ROS)产生、调节抗氧化酶基因表达以及增强抗氧化防护系统方面的作用,着重讨论了在ABA诱导的抗氧化防护过程中Ca2 、NADPH氧化酶与ROS之间的交谈机制.     

脱落酸与植物细胞的抗氧化防护

水分胁迫是一种影响植物生长发育、限制植物产量的重要胁迫因子。植物能够通过感知刺激、产生和传导信号、启动各种防护机制来响应与适应水分胁迫。植物激素脱落酸(ABA)作为一种胁迫信号,在调节植物对水分胁迫的反应中起着重要的作用。ABA不仅能诱导气孔关闭,而且能诱导编码耐脱水蛋白的基因表达。正在增加的证据显示,ABA增强水分胁迫的耐性与其诱导抗氧化防护系统有关。本文综述了ABA在诱导活性氧(ROS)产生、调节抗氧化酶基因表达以及增强抗氧化防护系统方面的作用,着重讨论了在ABA诱导的抗氧化防护过程中Ca2 、NADPH氧化酶与ROS之间的交谈机制。