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。     

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。     

不同土壤水分条件下土壤容重对玉米木质部汁液中ABA浓度和气孔导度的影响

受旱玉米植株木质部汁液中的ABA浓度是高水分处理的5倍,土壤容重每增加0.12g·cm-3,木质部液汁中ABA浓度约增加1倍。在相同土壤基质势下,植株的气孔导度和蒸腾速率随土壤容重的增大而下降,而容重对植株的叶水势没有影响。生长在混合容重(一边为1.20g·cm-3,另一边为1.45g·cm-3)土壤上的植株中,ABA浓度和气孔导度与全部根系处在高容重土壤中的植株接近。     

周期性土壤干旱和叶片水势对气孔响应木质部ABA灵敏度的影响

研究了周期性土壤干旱期间气孔对木质部ＡＢＡ响应的灵敏度的变化以及叶片水势对灵敏度的影响。实验结果证明了木质部ＡＢＡ浓度是反映根系周围土壤水分状况的一个指标的结论。土壤周期性干旱不影响木质部ＡＢＡ浓度对土壤水分状况的依赖关系,但显著地提高了气孔对木质部ＡＢＡ 响应的灵敏度。根据对实测数据的数学模拟结果显示,引起气孔导度下降５０％ 所需的木质部ＡＢＡ浓度从第一轮土壤干旱的７５０ ｎｍｏｌ／Ｌ降至第二轮土壤干旱的５５０ ｎｍｏｌ／Ｌ。分根实验的结果表明,叶片水分亏缺显著提高了气孔对木质部ＡＢＡ 的响应的灵敏程度,全根干旱中引起气孔导度下降５０ ％ 所需的木质部ＡＢＡ 浓度比半根干旱的小２ ～４ 倍。这表明,气孔对木质部ＡＢＡ响应的灵敏度不是一个固定的特性,可随植物生长环境及许多其他因素的变化而表现出很大的差异     

CO2浓度、氮和水分对春小麦光合、蒸散及水分利用效率的影响

研究了不同土壤氮和土壤水分条件下,大气CO2浓度升高对春小麦光合作用、气孔导度、蒸散和水分利用效率的影响。结果表明,CO2浓度升高,干旱处理的春小麦（Triticum aestivum L.）叶片光合作用速率幅度增加大于湿润处理,随着氮肥用量增加光合速率相应增加,而不施氮脂增加有限;干旱处理气孔导度幅度减少大于湿润处理,不施氮肥的大于氮肥充足的CO2浓度升高,干旱处理的蒸散量减少比湿润处理多,不施氮肥的蒸散量减少较为明显;但干旱处理单叶WUE增加大于湿润处理;随着氮肥用量增加,冠层WUE提高,而不施氮肥的冠层WUE最低。因而CO2浓度升高、光合速率增加和蒸散量减少会减缓干旱的不利影响,增强作物对干旱胁迫的抵御能力。     

干旱和复水对膜下滴灌棉花根系及叶片内源激素含量的影响

新疆气候生态条件下,采用膜下滴灌技术,在棉花不同生育时期设置不同程度干旱处理,研究干旱和复水对棉花根系及叶片内源激素含量和叶片气孔导度的影响.结果表明:在不同生育阶段,随土壤含水率的降低,根系及叶片脱落酸(ABA)含量显著增加,玉米素(ZRs)含量减少,叶片气孔导度(gs)和光合速率(Pn)显著降低,以初花-盛花期土壤相对含水率为40％ ～ 45％处理降幅较大.土壤干旱处理结束后复水,根系及叶片ABA含量并未随土壤水分条件的改善而降低,根系ZRs含量在复水后1～3d均可恢复或超过对照,与叶片gs呈正相关,其中以盛蕾-初花期土壤相对含水率为50％～55％处理棉株叶片ZRs含量和gs恢复速度快、强度大.说明干旱后复水根系较高的ZRs含量是导致其叶片gs和Pn较高的主要原因.     

根源ABA参与气孔调节的数学模拟

建立了包括植物体内的水分传输,并有根源ABA参与的气孔调节模型,模拟了饱和水气压差(VPD)、气温、表层土壤含水量(θ_(s1))等环境因子对叶片水势、木质部汁液中ABA浓度([ABA]_x)及气孔导度的影响。结果显示,VPD和气温的变化能够改变叶片水势及气孔导度;[ABA]_x几乎不受VPD和气温变化的影响,却决定着叶片水势及气孔导度对VPD和气温变化的响应幅度;θ_(s1)影响[ABA]_x,并由此影响气孔导度,但相比之下对叶片水势的作用并不显著。     

CO2浓度、氮和水分对春小麦光合、蒸散及水分利用效率的影响

研究了不同土壤氮和土壤水分条件下,大气CO₂浓度升高对春小麦光合作用、气孔导度、蒸散和水分利用效率的影响.结果表明,CO₂浓度升高,干旱处理的春小麦(Triticum aestivum L.)叶片光合作用速率幅度增加大于湿润处理,随着氮肥用量增加光合速率相应增加,而不施氮肥增加有限;干旱处理气孔导度幅度减少大于湿润处理,不施氮肥的大于氮肥充足的.CO₂浓度升高,干旱处理的蒸散量减少比湿润处理多,不施氮肥的蒸散量减少较为明显;但干旱处理单叶WUE增加大于湿润处理;随着氮肥用量增加,冠层WUE提高,而不施氮肥的冠层WUE最低.因而CO₂浓度升高、光合速率增加和蒸散量减少会减缓干旱的不利影响,增强作物对干旱胁迫的抵御能力.     

土壤水分亏缺条件下根源信号ABA参与作物气孔调控的数值模拟

建立了根系吸水模型和根源ABA参与作物气孔调控过程相耦合的气孔导度模型,该模型在根源信号ABA的产生项中考虑了根系吸水影响函数和根系密度分布函数.利用该耦合模型模拟大田状况下根源ABA参与玉米气孔行为调控过程,结果表明,由于充分考虑了根区土壤水势和土壤中根长密度分布对根系吸水的影响,较好地反映了土壤不同层次根系吸水强度,更为确切地描述了当土壤水分亏缺时,根系合成ABA的量、各层根系蒸腾流中ABA浓度、木质部ABA浓度以及最终ABA参与对气孔行为的调控作用.     

土壤干旱条件下玉米叶片内源激素含量及光合作用的变化

利用HPLC和ELISA技术研究了土壤干旱条件下玉米叶片内源IAA、ABA、ZR、DHZR、iPA含量的变化情况 ,以及叶片光合作用过程中 ,光合速率 (Pn) ,气孔导度 (Gs) ,PSⅡ光化学效率 (Fv/Fm)的变化情况 ,结果发现 :叶片中IAA浓度 ,在整个干旱过程中变化不大 ,与对照相比下降不明显 ;IAA的浓度对干旱的反应不敏感 ;叶片中ABA浓度在干旱最初 3d里急剧升高 ,直至最大值 ,之后有所下降 ;ABA的浓度对干旱反应敏感 ,但在整个干旱过程中ABA浓度并不随土壤相对含水量的减少而逐步升高 ;叶片中ZR和DHZR的浓度在干旱过程中与对照相比变化不明显 ,iPA浓度在干旱 4d后显著下降 ;叶片Pn在干旱初始 4d里随RWC的减小而缓慢下降 ,4d之后下降迅速 ;Gs从干旱第一天起即迅速减小 ,到第 4天近乎于零 ,Fv/Fm从干旱第 5天后开始逐步减小。这些结果证实在干旱过程中叶片ABA浓度的升高对气孔导度的调节作用 ,干旱初期光合速率的下降主要是气孔关闭所致 ;干旱后期光合速率的快速下降可能是PSⅡ光反应效率降低造成的     

Hydraulic and Chemical Responses of Citrus Seedlings to Drought and Osmotic Stress

In this work we investigated the function of abscisic acid (ABA) as a long-distance chemical signal communicating water shortage from the root to the shoot in citrus plants. Experiments indicated that stomatal conductance, transpiration rates, and leaf water potential decline progressively with drought. ABA content in roots, leaves, and xylem sap was also increased by the drought stress treatment three- to sevenfold. The addition of norflurazon, an inhibitor of ABA biosynthesis, significantly decreased the intensity of the responses and reduced ABA content in roots and xylem fluid, but not in leaves. Polyethylene glycol (PEG)-induced osmotic stress caused similar effects and, in general, was counteracted only by norflurazon at the lowest concentration (10%). Partial defoliation was able to diminish only leaf ABA content (22.5%) at the highest PEG concentration (30%), probably through a reduction of the active sites of biosynthesis. At least under moderate drought (3–6 days without irrigation), mechanisms other than leaf ABA concentration were required to explain stomatal closure in response to limited soil water supply. Measurements of xylem sap pH revealed a progressive alkalinization through the drought condition (6.4 vs. 7.1), that was not counteracted with the addition of norflurazon. Moreover, in vitro treatment of detached leaves with buffers iso-osmotically adjusted at pH 7.1 significantly decreased stomatal conductance (more than 30%) as much as 70% when supplemented with ABA. Taken together, our results suggest that increased pH generated in drought-stressed roots is transmitted by the xylem sap to the leaves, triggering reductions in shoot water loss. The parallel rise in ABA concentration may act synergistically with pH alkalinization in xylem sap, with an initial response generated from the roots and further promotion by the stressed leaves.     

Does engineering abscisic acid biosynthesis in Nicotiana plumbaginifolia modify stomatal response to drought?

The consequences of manipulating abscisic acid (ABA) biosynthesis rates on stomatal response to drought were analysed in wild‐type, a full‐deficient mutant and four under‐producing transgenic lines of N. plumbaginifolia. The roles of ABA, xylem sap pH and leaf water potential were investigated under four experimental conditions: feeding detached leaves with varying ABA concentration; injecting exogenous ABA into well‐watered plants; and withholding irrigation on pot‐grown plants, either intact or grafted onto tobacco. Changes in ABA synthesis abilities among lines did not affect stomatal sensitivity to ABA concentration in the leaf xylem sap ([ABA]_xyl), as evidenced with exogenous ABA supplies and natural increases of [ABA]_xyl in grafted plants subjected to drought. The ABA‐deficient mutant, which is uncultivable under normal evaporative demand, was grafted onto tobacco stock and then presented the same stomatal response to [ABA]_xyl as wild‐type and other lines. This reinforces the dominant role of ABA in controlling stomatal response to drought in N. plumbaginifolia whereas roles of leaf water potential and xylem sap pH were excluded under all studied conditions. However, when plants were submitted to soil drying onto their own roots, stomatal response to [ABA]_xyl slightly differed among lines. It is suggested, consistently with all the results, that an additional root signal of soil drying modulates stomatal response to [ABA]_xyl.     

Maize stomatal conductance in the field: its relationship with soil and plant water potentials, mechanical constraints and ABA concentration in the xylem sap

Abstract. Stomatal conductance, leaf water potential, soil water potential and concentration of abscisic acid (ABA) in the xylem sap were measured on maize plants growing in the field, in two treatments with contrasting soil structures. Soil compaction affected the stomatal conductance, but this effect was no longer observed if the soil water potential was increased by irrigation. Differences in leaf water potential did not account for the differences in conductance between treatments. Conversely, the relationship between stomatal conductance and concentration of ABA in the xylem sap was consistent during the experiment. The proposed interpretation is that stomatal conductance was controlled by the root water potential via an ABA message. Control of the stomatal conductance by the leaf water potential or by an effect of mechanical stress on the roots is unlikely.     

Xylem ABA controls the stomatal conductance of field-grown maize subjected to soil compaction or soil drying

Stomatal conductance of individual leaves was measured in a maize field, together with leaf water potential, leaf turgor, xylem ABA concentration and leaf ABA concentration in the same leaves. Stomatal conductance showed a tight relationship with xylem ABA, but not with the current leaf water status or with the concentration of ABA in the bulk leaf. The relationship between stomatal conductance and xylem [ABA] was common for variations in xylem [ABA] linked to the decline with time of the soil water reserve, to simultaneous differences between plants grown on compacted, non-compacted and irrigated soil, and to plant-to-plant variability. Therefore, this relationship is unlikely to be fortuitous or due to synchronous variations. These results suggest that increased concentration of ABA in the xylem sap in response to stress can control the gas exchange of plants under field conditions.     

Drought-induced changes in xylem pH, ionic composition, and ABA concentration act as early signals in field-grown maize (Zea mays L.).

Early signals potentially regulating leaf growth and stomatal aperture in field-grown maize (Zea mays L.) subjected to drought were investigated. Plants grown in a field lysimeter on two soil types were subjected to progressive drought during vegetative growth. Leaf ABA content, water status, extension rate, conductance, photosynthesis, nitrogen content, and xylem sap composition were measured daily. Maize responded similarly to progressive drought on both soil types. Effects on loam were less pronounced than on sand. Relative to fully-watered controls, xylem pH increased by about 0.2 units one day after withholding irrigation (DAWI) and conductivity decreased by about 0.25 mS cm(-1) 1-3 DAWI. Xylem nitrate, ammonium, and phosphate concentrations decreased by about 50% at 1-5 DAWI and potassium concentration decreased by about 50% at 7-8 DAWI. Xylem ABA concentration consistently increased by 45-70 pmol ml(-1) at 7 DAWI. Leaf extension rate decreased 5 DAWI, after the changes in xylem chemical composition had occurred. Leaf nitrogen significantly decreased 8-16 DAWI in droughted plants. Midday leaf water potential and photosynthesis were significantly decreased in droughted plants late in the drying period. Xylem nitrate concentration was the only ionic xylem sap component significantly correlated to increasing soil moisture deficit and decreasing leaf nitrogen concentration. Predawn leaf ABA content in droughted plants increased by 100-200 ng g(-1) dry weight at 7 DAWI coinciding with a decrease in stomatal conductance before any significant decrease in midday leaf water potential was observed. Based on the observed sequence, a chain of signal events is suggested eventually leading to stomatal closure and leaf surface reduction through interactive effects of reduced nitrogen supply and plant growth regulators under drought.     

Does Xylem Sap ABA Control the Stomatal Behaviour of Water-Stressed Sycamore (Acer pseudoplatanus L.) Seedlings?

Sycamore seedlings were grown with their root systems dividedequally between two containers. Water was withheld from onecontainer while the other container was kept well-watered. Effectsof soil drying on stomatal behaviour, shoot water status, andabscisic acid (ABA) concentration in roots, xylem sap and leaveswere evaluated. At 3 d, root ABA in the drying container increased significantly,while the root ABA in the unstressed container of the same plantsdid not differ from that of the control. The increase in rootABA was associated with the increase in xylem sap ABA and withthe decrease in stomatal conductance without any significantperturbation in shoot water status. At 7 d, despite the continuous increase in root ABA concentration,xylem sap ABA showed a marked decline when soil water contentwas depleted below 013 g g^–1. This reduction in xylemsap ABA coincided with a partial recovery of stomatal conductance.The results indicate that xylem sap ABA is a function of rootABA as well as the flow rate of water from roots to shoots,and that this ABA can be a sensitive indicator to the shootof the effect of soil drying. Key words: Acer pseudoplatanus L., soil drying, stomatal behaviour, xylem sap ABA     

Stomatal conductance and xylem sap properties of aspen (Populus tremuloides) in response to low soil temperature

The mechanisms regulating stomatal response following exposure to low (5°C) soil temperature were investigated in aspen ( Populus tremuloides Michx.) seedlings. Low soil temperature reduced stomatal conductance within 4 h, but did not alter shoot xylem pressure potential within 24 h. The xylem sap composition was altered and its pH increased from 6.5 to 7.1 within the initial 4 h of the low temperature treatment. However, the increase in abscisic acid (ABA) concentration in xylem sap was observed later, after 8 h of treatment. These changes were accompanied by a reduction in the electrical conductivity and an increase in the osmotic potential of the xylem sap. The timing of physiological responses to low soil temperature suggests that the rapid pH change of the xylem sap and accompanying changes in ion concentration were the initial factors which triggered stomatal closure in low temperature-treated seedlings, and that the role of the more slowly accumulating ABA was likely to reinforce the stomatal closure. When leaf discs were exposed to xylem sap extracted from low soil temperature-treated plants, stomatal aperture was negatively correlated with ABA and positively correlated with K⁺ concentrations of the xylem sap. The stomatal opening in the leaf discs linearly increased in response to exogenous KCl concentrations when K⁺ concentrations were in the similar range to those detected in the xylem sap. The lowest concentration of exogenous ABA to induce stomatal closure was several-fold higher compared with the concentration present in the xylem sap.     

How the roots contribute to the ability of Phaseolus vulgaris L. to cope with chilling-induced water stress

Intact plants and stem-girdled plants of Phaseolus vulgaris grown hydroponically were exposed to 5 degrees C for up to 4 d; stem girdling was used to inhibit the phloem transport from the leaves to the roots. After initial water stress, stomatal closure and an amelioration of root water transport properties allowed the plants to rehydrate and regain turgor. Chilling augmented the concentration of abscisic acid (ABA) content in leaves, roots and xylem sap. In intact plants stomatal closure and leaf ABA accumulation were preceded by a slight alkalinization of xylem sap, but they occurred earlier than any increase in xylem ABA concentration could be detected. Stem girdling did not affect the influence of chilling on plant water relations and leaf ABA content, but it reduced slightly the alkalinization of xylem sap and, principally, prevented the massive ABA accumulation in root tissues and the associated transport in the xylem that was observed in non-girdled plants. When the plants were defoliated just prior to chilling or after 10 h at 5 degrees C, root and xylem sap ABA concentration remained unchanged throughout the whole stress period. When the plants were chilled under conditions preventing the occurrence of leaf water deficit (i.e. at 100% relative humidity), there were no significant variations in endogenous ABA levels. The increase in root hydraulic conductance in chilled plants was a response neither to root ABA accretion, nor to some leaf-borne chemical signal transported downwards in the phloem, nor to low temperature per se, as indicated by the results of the experiments with defoliated or girdled plants and with plants chilled at 100% relative humidity. It was concluded that the root system contributed substantially to the bean's ability to cope with chilling-induced water stress, but not in an ABA-dependent manner.     

Changes in the concentration of ABA in xylem sap as a function of changing soil water status can account for changes in leaf conductance and growth

Abstract. Maize seedlings ( Zea mays L. John Innes F1 hybrid) were grown in a greenhouse in l-m-long tubes of soil. When the plants were well established, water was withheld from half of the tubes. Control plants were watered every day during the 20-d experimental period. The soil drying treatment resulted in a substantial restriction of stomatal conductance and a limitation in shoot growth, even though there was no detectable difference in the water relations of watered and unwatered plants. From day 7 of the soil drying treatment, xylem ABA concentrations (measured using the sap exuded from detopped plants) were substantially increased in unwatered plants compared to values recorded with sap from plants watered every day. Measurements of water potential through the profile of unwatered soil suggest that xylem ABA concentrations reflects the extent of soil drying. Leaf ABA content was a much less sensitive indicator of the effect of soil drying and during the whole of experimental period there was no significant difference between ABA concentration in leaves of well watered and unwatered plants. In a second set of experiments, ABA was fed to part of the roots of potted maize plants to manipulate xylem ABA concentration. These manipulations suggested that the increases in ABA concentration in xylem sap, which resulted from soil drying, were adequate to explain the observed variation in stomatal conductance and might also explain the restriction in leaf growth rate. These results are discussed in the light of recent work which suggests that stomatal responses to soil drying are partly attributable to an as-yet unidentified inhibitor of stomatal opening.     

Long-distance ABA Signaling and Its Relation to Other Signaling Pathways in the Detection of Soil Drying and the Mediation of the Plant’s Response to Drought

In this article we review evidence for a variety of long-distance signaling pathways involving hormones and nutrient ions moving in the xylem sap. We argue that ABA has a central role to play, at least in root-to-shoot drought stress signaling and the regulation of functioning, growth, and development of plants in drying soil. We also stress the importance of changes in the pH of the leaf cell apoplast as influenced both by edaphic and climatic variation, as a regulator of shoot growth and functioning, and we show how changes in xylem and apoplastic pH can affect the way in which ABA regulates stomatal behavior and growth. The sensitivity to drought of the pH/ABA sensing and signaling mechanism is emphasized. This allows regulation of plant growth, development and functioning, and particularly shoot water status, as distinct from stress lesions in growth and other processes as a reaction to perturbations such as soil drying.