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。     

土壤干旱条件下氮素营养对玉米内源激素含量影响

在田间持水量分别保持于35％、55％和75％±5％的土壤水分条件下,利用盆栽实验研究了土壤干旱和氮素营养对玉米内源激素和气孔导度的影响．结果表明,土壤干旱下氮素营养明显降低了玉米根系木质部汁液ABA浓度,而正常供水下施氮处理间则无显著差异(施氮处理仍较低),同时测定的叶片ABA浓度则呈相反的变化趋势,表现为干旱下施氮处理要高于不施氮处理;施氮处理木质部汁液中ZRs浓度应低于相应的不施氮处理,在调控气孔行为方面并未表现拮抗ABA作用;3种土壤水分条件下,施氮玉米叶片的气孔导度均高于不施氮处理,与木质部汁液ABA浓度呈负相关,说明施氮处理较低的根源ABA浓度是导致其气孔导度较大的主要原因．     

Multiple signals and mechanisms that regulate leaf growth and stomatal behaviour during water deficit

We highlight the novel observation that the responses of maize leaf growth to abscisic acid (ABA) signals can be amplified both by mild water deficits and by nutrient stress. Under our experimental conditions these stresses alone had no effect on leaf growth rate. In most cases leaf growth responses were not attributable to changes in the turgors of growing cells, focusing attention on a regulatory role for cell wall biochemistry. Roles for xyloglucan endotransglycosylase (XET), expansins and peroxidases are discussed. An effect of drought on the activity of expansins seems particularly attractive if xylem sap pH is considered as a chemical signal. We show how changes in xylem sap pH can also modify local accumulation of ABA and thereby modify the apparent sensitivity of guard cells to ABA signals.     

不同水分条件下两个树种木质部栓塞对P素添加的响应

 在两种水分供给(干旱胁迫和适宜水分,土壤含水量分别为田间持水量的30%～40%和70%～80%)下,研究了耐旱树种元宝枫(Acer truncatum)和 中生树种女贞(Ligustrum lucidum )木质部栓塞(以导水率(Percentage loss of hydraulic conductivity, PLC)损失程度衡量)对P素添加的 响应。结果发现,两个树种PLC的日变化均呈现出先上升后降低的规律,表明木质部栓塞的形成与恢复是植物体的一种平常事件;除适宜水分条 件的女贞外,P素可以显著提高元宝枫和遭受干旱胁迫时女贞的PLC;两种水分条件下,干旱胁迫时元宝枫木质部栓塞明显高于适宜水分供给时 。女贞的PLC在两种水分状况下无显著差异;树种间,干旱胁迫促进了元宝枫木质部的栓塞形成,明显高于同等水分条件下的女贞。该研究结果 证实了“木质部限流耐旱假设”。     

Rapid responses of C14 clone of Eucalyptus globulus to root drought stress: Time-course of hormonal and physiological signaling

The responses of juvenile plants of forest crops to drought stress are a key stage in the survival of forest populations. In this work, a suitable experimental system to study the early drought resistance mechanisms and signaling in a drought-tolerant clone (C14) of Eucalyptus globulus Labill is proposed. This system, using hydroponic culture and an osmotic agent, polyethylene glycol 8000, was demonstrated to induce severe stress in the root area, affecting the responses of the plantlets at the aerial level. These responses were very fast, beginning only 3 h after the induction of stress, and the results highlight the roles of xylematic abscisic acid (ABA) and pH changes over other signals, such as cytokinins, as early chemical signals in rapid water stress. The relationship between these chemical factors, ABA and pH, and the physiological and water parameters observed were significant, supporting their proposed principal role. This work aids our understanding of underlying responses to hydrological limitations of forest crops, and provides valuable information for further physiological and molecular studies of water stress in this and other tree species.     

Reproductive water supply is prioritized during drought in tomato

Reproductive success largely defines the fitness of plant species. Understanding how heat and drought affect plant reproduction is thus key to predicting future plant fitness under rising global temperatures. Recent work suggests reproductive tissues are highly vulnerable to water stress in perennial plants where reproductive sacrifice could preserve plant survival. However, most crop species are annuals where such a strategy would theoretically reduce fitness. We examined the reproductive strategy of tomato (Solanum lycopersicum var. Rheinlands Ruhm) to determine whether water supply to fruits is prioritized above vegetative tissues during drought. Using optical methods, we mapped xylem cavitation and tissue shrinkage in vegetative and reproductive organs during dehydration to determine the priority of water flow under acute water stress. Stems and peduncles of tomato showed significantly greater xylem cavitation resistance than leaves. This maintenance of intact water supply enabled tomato fruit to continue to expand during acute water stress, utilizing xylem water made available by tissue collapse and early cavitation of leaves. Here, tomato plants prioritize water supply to reproductive tissues, maintaining fruit development under drought conditions. These results emphasize the critical role of water transport in shaping life history and suggest a broad relevance of hydraulic prioritization in plant ecology.     

Stomatal factors and vulnerability of stem xylem to cavitation in poplars

The relationships between the vulnerability of stem xylem to cavitation, stomatal conductance, stomatal density, and leaf and stem water potential were examined in six hybrid poplar (P38P38, Walker, Okanese, Northwest, Assiniboine and Berlin) and balsam poplar (Populus balsamifera) clones. Stem xylem cavitation resistance was examined with the Cavitron technique in well-watered plants grown in the greenhouse. To investigate stomatal responses to drought, plants were subjected to drought stress by withholding watering for 5 (mild drought) and 7 (severe drought) days and to stress recovery by rewatering severely stressed plants for 30 min and 2 days. The clones varied in stomatal sensitivity to drought and vulnerability to stem xylem cavitation. P38P38 reduced stomatal conductance in response to mild stress while the balsam poplar clone maintained high leaf stomatal conductance under more severe drought stress conditions. Differences between the severely stressed clones were also observed in leaf water potentials with no or relatively small decreases in Assiniboine, P38P38, Okanese and Walker. Vulnerability to drought-induced stem xylem embolism revealed that balsam poplar and Northwest clones reached loss of conductivity at lower stem water potentials compared with the remaining clones. There was a strong link between stem xylem resistance to cavitation and stomatal responsiveness to drought stress in balsam poplar and P38P38. However, the differences in stomatal responsiveness to mild drought suggest that other drought-resistant strategies may also play a key role in some clones of poplars exposed to drought stress.     

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.     

Metabolomic and proteomic changes in the xylem sap of maize under drought

Plants produce compounds in roots that are transported to shoots via the xylem sap. Some of these compounds are vital for signalling and adaptation to environmental stress such as drought. In this study, we screened the xylem sap using mass spectrometry to quantify the changes in new and previously identified sap constituents under extended drought. We detected and quantified the changes in the concentration of 31 compounds present in the xylem sap under progressively increasing drought stress. We found changes in the hormones abscisic acid (ABA) and cytokinin, and the presence of high concentrations of the aromatic cytokinin 6-benzylaminopurine (BAP). Several phenylpropanoid compounds (coumaric, caffeic and ferulic acids) were found in xylem sap. The concentrations of some of these phenylpropanoid compounds changed under drought. In parallel, an analysis of the xylem sap proteome was conducted. We found a higher abundance of cationic peroxidases, which with the increase in phenylpropanoids may lead to a reduction in lignin biosynthesis in the xylem vessels and could induce cell wall stiffening. The application of new methodologies provides insights into the range of compounds in sap and how alterations in composition may lead to changes in development and signalling during adaptation to drought.     

Long-distance stress and developmental signals associated with abscisic acid signaling in environmental responses

Flowering plants consist of highly differentiated organs, including roots, leaves, shoots and flowers, which have specific roles: root system for water and nutrient uptake, leaves for photosynthesis and gas exchange and reproductive organs for seed production. The communication between organs through the vascular system, by which water, nutrient and signaling molecules are transported, is essential for coordinated growth and development of the whole plant, particularly under adverse conditions. Here, we highlight recent progress in understanding how signaling pathways of plant hormones are associated with long-distance stress and developmental signals, with particular focus on environmental stress responses. In addition to the root-to-shoot peptide signal that induces abscisic acid accumulation in leaves under drought stress conditions, we summarize the diverse stress-responsive peptide signals reported to date to play a role in environmental responses.     

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.     

Priming xylem for stress recovery depends on coordinated activity of sugar metabolic pathways and changes in xylem sap pH

Some plant species are capable of significant reduction of xylem embolism during recovery from drought despite stem water potential remains negative. However, the functional biology underlying this process is elusive. We subjected poplar trees to drought stress followed by a period of recovery. Water potential, hydraulic conductivity, gas exchange, xylem sap pH, and carbohydrate content in sap and woody stems were monitored in combination with an analysis of carbohydrate metabolism, enzyme activity, and expression of genes involved in sugar metabolic and transport pathways. Drought resulted in an alteration of differential partitioning between starch and soluble sugars. Upon stress, an increase in the starch degradation rate and the overexpression of sugar symporter genes promoted the efflux of disaccharides (mostly maltose and sucrose) to the apoplast. In turn, the efflux activity of the sugar‐proton cotransporters caused a drop in xylem pH. The newly acidic environment induced the activity of apoplastic invertases leading to the accumulation of monosaccharides in the apoplast, thus providing the main osmoticum necessary for recovery. During drought and recovery, a complex network of coordinated molecular and biochemical signals was activated at the interface between xylem and parenchyma cells that appeared to prime the xylem for hydraulic recovery.     

Cavitation resistance of peduncle,petiole and stem is correlated with bordered pit dimensions in Magnolia grandiflora

Variation in resistance of xylem to embolism among flowers, leaves, and stems strongly influences the survival and reproduction of plants. However, little is known about the vulnerability to xylem embolism under drought stress and their relationships to the anatomical traits of pits among reproductive and vegetative organs. In this study, we investigated the variation in xylem vulnerability to embolism in peduncles, petioles, and stems in a woody plant, Magnolia grandiflora. We analyzed the relationships between water potentials that induced 50% embolism (P₅₀) in peduncles, petioles, and stems and the conduit pit traits hypothesized to influence cavitation resistance. We found that peduncles were more vulnerable to cavitation than petioles and stems, supporting the hypothesis of hydraulic vulnerability segmentation that leaves and stems are prioritized over flowers during drought stress. Moreover, P₅₀ was significantly correlated with variation in the dimensions of inter-vessel pit apertures among peduncles, petioles and stems. These findings highlight that measuring xylem vulnerability to embolism in reproductive organs is essential for understanding the effect of drought on plant reproductive success and mortality under drought stress.     

Signalling mechanisms involved in the response of two varieties of Humulus lupulus L. to soil drying: II. changes in the concentration of abscisic acid catabolites and stress-induced phytohormones

Abscisic acid (ABA) is one of the most common stress signals that appear in plant organs in response to soil drying. Equilibrium between ABA biosynthesis and catabolism regulates ABA accumulation in plants under water stress. The aim of our work was to explore the dynamics of changes in ABA metabolites as well as other stress-induced phytohormones such as jasmonic acid, indole-3-acetic acid, and their respective metabolites in hop [Humulus lupulus (L.)] plants during drying and to identify among them potential signals involved in drought signalling. We showed that the concentrations of all ABA metabolites (except the concentration of ABA glucosyl ester in leaves) increased in the same manner in leaves and xylem sap approximately at the same level of soil water content when the relative water content of leaves decreased. The predominant metabolites in leaves and xylem sap were phaseic acid and dihydroxyphaseic acid. ABA glucosyl ester was not a source of the increased concentration of ABA in leaves and xylem sap because of its considerably lower concentration compared to ABA. The concentration of jasmonates decreased in leaves of hop plants. Changes in auxin concentration suggest that this hormone is involved in the response of hop plants to soil drying.     

Irrigation effects on daily and seasonal variations of trunk sap flow and leaf water relations in olive trees

Irrigation effects on whole-plant sap flow and leaf-level water relations were characterised throughout a growing season in an experimental olive (Olea europaea L.) orchard. Atmospheric evaporative demand and soil moisture conditions for irrigated and non-irrigated olive trees were also monitored. Whole-plant water use in field-grown irrigated and rain fed olive trees was determined using a xylem sap flow method (compensation heat-pulse velocity). Foliage gas exchange and water potentials were determined throughout the experimental period. Physiological parameters responded diurnally and seasonally to variations in tree water status, soil moisture conditions and atmospheric evaporative demand. There was a considerable degree of agreement between daily transpiration deduced from heat-pulse velocity and that determined by calibration using the Penman–Monteith equation in the field. Summer drought caused decreasing leaf gas exchange and water potentials, and a progressive increase in hydraulic conductance (stronger in non-irrigated than irrigated trees), probably attributable to modifications in hydraulic properties at the soil-root interface. Negligible hysteresis, attributable to low plant capacitance, was observed in the relationship between leaf water potential and sap flow. A proportional decrease in maximum daily leaf conductance with increasing vapour pressure deficit was observed, while mean daytime canopy stomatal conductance decreased with the season. As a result, plant water use was limited and excessive drought stress prevented. Non-irrigated olive trees recovered after the summer drought, showing a physiological behaviour similar to that of irrigated trees. In addition to physiological and environmental factors, there are endogenous keys (chemical signals) influencing leaf level parameters. Olive trees are confirmed to be economical and sparing users of soil water, with an efficient xylem sap transport, maintenance of significant gas exchange and transpiration, even during drought stress.     

Hydraulic and chemical signalling in the control of stomatal conductance and transpiration.

Abscisic acid (ABA) transported in the xylem from root to shoot and perceived at the guard cell is now widely studied as an essential regulating factor in stomatal closure under drought stress. This provides the plant with a stomatal response mechanism in which water potential is perceived in the root as an indication of soil water status and available water resources. There is also ample evidence that stomata respond directly to some component of leaf water status. This provides additional information about water potential gradients developing between root and shoot as the result of water transport, allowing for a more stable regulation of shoot water status and better protection of the transport system itself. The precise location at which leaf water status is sensed, however, and the molecular events transducing this signal into a guard cell response are not yet known. Major questions therefore remain unanswered on how water stress signals perceived at root and leaf locations are integrated at the guard cell to control stomatal behaviour.     

荒漠河岸林植物木质部导水与栓塞特征及其对干旱胁迫的响应

 以同处于干旱区的塔里木河下游(铁干里克)和黑河下游(乌兰图格)断面为研究区, 比较了荒漠河岸林主要建群种胡杨(Populus euphratica)、柽柳(Tamarix spp.)、疏叶骆驼刺(Alhagi sparsifolia)和花花柴(Karelinia caspia)在长期遭受不同干旱胁迫下的根、枝条木质部导水力和栓塞化程度的变化特征, 并分析了木质部导水对干旱胁迫的响应及适应策略。结果表明: 1) 黑河下游荒漠河岸林植物的导水能力显著高于塔里木河下游, 其中柽柳、胡杨、疏叶骆驼刺和花花柴根木质部的初始比导率(K_s0)分别高11.97、6.74、7.10和3.73倍, 枝条的K_s0分别高9.48、3.65、2.07和1.88倍, 地下水埋深导致的干旱胁迫程度不同是诱发荒漠植物导水能力差异的根本原因; 2)柽柳耐干旱能力最强, 适应范围较宽, 而花花柴、疏叶骆驼刺的耐旱性相对较弱, 适生范围较窄, 这可能与植物的根系分布有关; 3)干旱胁迫较轻时, 枝条木质部是荒漠河岸林植物水分传输的主要阻力部位, 干旱胁迫严重时, 根木质部是限制植株水流的最大阻碍部位; 4)荒漠河岸林植物主要通过调节枝条木质部的水流阻力来适应干旱胁迫, 且其适应策略与干旱胁迫程度有关, 干旱胁迫轻时, 植物通过限制枝条木质部水流来协调整株植物的均匀生长; 干旱胁迫严重时, 植物通过牺牲劣势枝条、增强优势枝条水流来提高植株整体生存的机会。     

Anatomical adaptations of Astragalus gombiformis Pomel. under drought stress

The present study was designed to study the effect of drought on root, stem and leaf anatomy of Astragalus gombiformis Pomel. Several root, stem and leaf anatomical parameters (cross section diameter, cortex, root cortical cells, pith, leaf lamina and mesophyll thickness) were reduced under moderate to severe water deficit (20–30 days of withheld irrigation). The stele/cross section root ratio increased under moderate water deficit. The root’s and stems vascular systems showed reduced xylem vessel diameter and increased wall thickness under water deficit. In addition, the root xylem vessel density was increased in these drought conditions while it was unchanged in the stems. The stomata density was increased under prolonged drought conditions whereas the stomata size was untouched. The leaf vascular system showed reduced xylem and phloem tissue thickness in the main vein under moderate to severe water deficit. However, in the lamina the vascular tissue and the distance between vascular bundle were unaffected. Our findings suggest a complex network of anatomical adaptations such as a reduced vessel size with increased wall thickness, lesser cortical and mesophyll parenchyma formation and increased stomata density. These proprieties are required for the maintenance of water potential and energy storage under water stress which can improve the resistance of A. gombiformis to survive in arid areas.     

植物木质部栓塞测定技术的研究进展

气候变化引发干旱频度和强度的变化影响植物的生长发育和生态适应。干旱胁迫会引起木质部栓塞并造成水力失效,而如何准确量化木质部抗栓塞的能力对研究植物对干旱的响应过程尤为重要。通常可通过脆弱性曲线量化木质部抗栓塞的能力。目前已经研发出构建木质部栓塞脆弱性曲线的多种方法,但不同方法往往产生不一致的结果。深入理解这些方法的设计原理并在实际应用时比较各方法的异同,对合理解释相关文献数据及准确选择干旱预测模型等尤为重要。本文阐述了自然干燥法、离心法、注气法、声学测定法、同步加速器与X射线显微断层扫描法、光学可视化法及抽气法7种测定木质部栓塞脆弱性的方法,并总结了近年来各测定方法在具体研究中的运用情况及存在的争议。最后,对未来研究测定木质部栓塞脆弱性与实际运用相关方法的选择等提出了展望。