Sequential response of whole plant water relations to prolonged soil drying and the involvement of xylem sap ABA in the regulation of stomatal behaviour of sunflower plants

Sunflower plants ( Helianihus animus cv. Tall Single Yellow} were grown in the greenhouse in drain pipes (100 mm inside diameter and 1 m long) rilled with John Innes No. 2 compost. When the fifth leaf had emerged, half of the plants were left unwatered for 6 days, rewatered for 2 days and then not watered for another 12 days. Measurements of water relations and abaxial stomatal conductance were made at each leaf position at regular intervals during the experimental period. Estimates were also made of soil water potentials along the soil profile and of ABA concentrations in xylem sap and leaves.
Soil drying led to some reduction in stomatal conductance alter only 3 days but leaf turgors were not reduced until day 13 (6 days after rewatering). When the water relations of leaves did change, older leases became substantially dehydrated while high turgors were recorded in younger leaves. Leaf ABA content measured on the third youngest leaf hardly changed over the first 13 days of the experiment, despite substantial soil drying, while xylem ABA concentrations changed very significantly and dynamically as soil water status varied, even when there was no effect of soil drying on leaf water relations. We argue that the highest ABA concentrations in the xylem, found as a result of substantial soil drying, arise from synthesis in both the roots and the older leaves, and act to delay the development of water deficit in younger leases.
In other experiments ABA solutions were watered on to the root systems of sunflower plants to increase ABA concentrations in xylem sap. The stomatal response to applied ABA was quantitatively very similar to that to ABA generated as a result of soil drying. There was a log-linear relationship between the reduction of leaf conductance and the increase of ABA concentration m xylem sap.     

The Effects of Partially Drying Part of the Root System of Helianthus annuus on the Abscisic Acid Content of the Roots, Xylem Sap and Leaves

Plants of Helianthus annuus were grown in soil in pots suchthat approximately 30% of the root system protruded throughthe base of the pot. After 7 d further growth in aerated nutrientsolution, the attached, protruding roots were air-dried for10–15 min and thereafter surrounded with moist still air,in the dark, for 49 h, whilst the soil was kept at field capacity.The roots of the control plants remained in the nutrient solutionthroughout the experiment. This treatment rapidly reduced the water content of protrudingroots from 20.5 to 17.8 g g^–1 dry mass (DM), which remainedless than that of the control roots for the rest of the experiment.This treatment also reduced root turgor and water potential.The abscisic acid (ABA) concentrations in the protruding roots,xylem sap and leaves of the treated plants increased significantly,compared to values recorded for control plants. In treated roots, the ABA concentration was significantly increased4 h after treatment, with a maximum of 4.4+0.1 nmol g^–1(DM) after 25 h. The ABA concentration in the xylem sap of thetreated plants was significantly greater than in the controls25 h, 30 h, and 49 h after the partial drying of the roots,with a maximum concentration of approximately 970 pmol ABA cm-3at 49 h. Initially, the ABA concentration in the leaves was0.45 nmol g^–1 (DM) which increased significantly to 1.1±0.1 nmol g^–1 at 25 h, to 1.7±0.3 nmol g^–1at 49 h. Leaf conductance was significantly less in plants with air-driedroots than in the controls 8 h after the start of the treatmentand thereafter. The water relations of the leaves of the treatedplants did not differ from those of the control plants. These results confirm previous reports that ABA is rapidly generatedin partially-dried and attached root systems and demonstratesa concomitant large increase in the ABA content of the xylemsap. It is suggested that partial dehydration of some of theroots of Helianthus annuus, increases ABA concentration in thetranspiration stream and decreases leaf conductance in the absenceof changes in leaf water status. As these responses were initiatedin free-growing roots the stimulus is independent of any increasesin soil shear strength that are associated with soil drying. Key words: Soil drying, roots, ABA, leaf conductance, water relations     

The relations of stomatal closure and reopening to xylem ABA concentration and leaf water potential during soil drying and rewatering

Two tropical tree species, Acacia confusa and Leucaena leucocephala, were used to study the relationships among stomatal conductance, xylem ABA concentration and leaf water potential during a soil drying and rewatering cycle. Stomatal conductance of both A. confusa and L. leucocephala steadily decreased with the decreases in soil water content and pre-dawn leaf water potential. Upon rewatering, soil water content and pre-dawn leaf water potential rapidly returned to the control levels, whereas the reopening of stomata showed an obvious lag time. The length of this lag time was highly dependent not only upon the degree of water stress but also on plant species. The more severe the water stress, the longer the lag time. When A. confusa and L. leucocephala plants were exposed to the same degree of water stress (around –2.0 MPa in pre-dawn leaf water potential), the stomata of A. confusa reopened to the control level 6 days after rewatering. However, it took L. leucocephala about 14 days to reopen fully. A very similar response of leaf photosynthesis to soil water deficit was also observed for both species. Soil drying resulted in a significant increase in leaf and xylem ABA concentrations in both species. The more severe the water stress, the higher the leaf and xylem ABA concentrations. Both leaf ABA and xylem ABA returned to the control level following relief from water deficit and preceded the full recovery of stomata, suggesting that the lag phase of stomatal reopening was not controlled by leaf and/or xylem ABA. In contrast to drying the whole root system, drying half of the root system did not change the leaf water relations, but caused a significant increase in xylem ABA concentration, which could fully explain the decrease of stomatal conductance. After rewatering, the stomatal conductance of plants in which half of the roots were dried recovered more rapidly than those of whole-root dried plants, indicating that the leaf water deficit that occurred during the drying period was related to the post-stress stomatal inhibition. These results indicated that the decrease in stomatal conductance caused by water deficit was closely related to the increase in xylem ABA, but xylem ABA could not fully explain the reopening of stomata after relief of water stress, neither did the leaf ABA. Some unknown physiological and/or morphological processes in the guard cells may be related to the recovery process.     

The role of abscisic acid and water relations in drought responses of subterranean clover

The role of water relations and abscisic acid (ABA) in the responsesto drought were studied in a mediterranean forage crop, Trifoliumsubterraneum L. under field conditions. Soil and plant waterstatus, leaf gas exchange parameters, and xylem sap ABA contentwere determined at different times during a long-term soil dryingepisode in irrigated and droughted plants. The diurnal time-coursesof these parameters were also measured at the end of a droughtperiod. In response to soil drying stomatal conductance (g) was reducedearly to 50% that of irrigated plants before any substantialchange in water potential was detected. A close logarithmicregression between photosynthesis rate (A) and g was present.For the first weeks of drought the decline in A was less pronouncedthan in g, thus increasing water use efficiency. Stomatal conductanceduring diurnal time-courses showed no consistent relationshipswith respect to etther ABA or leaf water potential. Throughoutthe experimental period dependence of g on leaf water statuswas evident from the tight correlation (r²=0.88, P<0.01)achieved between stomatal conductance and midday water potential,but the correlation was also high when comparing g with respectto ABA content in xylem sap (r=0.83, P<0.001). However, thestomata from drought acclimated plants were apparently moresensitive to xylem ABA content. For similar xylem ABA concentrationsstomatal conductance was significantly higher in irrigated thanin waterstressed plants. Key words: Drought, stomatal conductance, water potential, abscisic acid     

Control of crops leaf growth by chemical and hydraulic influences

Three species of forage grasses (Festuca arundinacea, Eragrostiscurvula, Sporobolus stapfianus) commonly grown in the Mediterraneanregion were subjected to a soil drying treatment. Leaf growthrate in F. arundinacea was highly sensitive to soil drying andlow growth rates were associated with high laminar turgors.The production of ABA was stimulated by soil drying and therewas a clear relation between increasing ABA accumulation andreduction in leaf growth. Leaf growth of E. cutvula, a C⁴ warmseason grass, was relatively insensitive to soil drying whichwas not accompanied by a substantial increase in leaf ABA content.S. stapfianus, a resurrection plant, was highly sensitive todecreasing soil water availability. In these two latter species,leaf growth was substantially restricted before ABA accumulationoccurred. It is suggested that reductions in laminar turgorof E. curvula and S. stapfianus may be limiting leaf growthas soil dries. The results indicated a different mechanism ofsensing and responding to reduction in soil water availabilityfor the three species studied. The relative importance of thechemical and hydraulic control of leaf growth is discussed. Key words: Leaf growth, water relations, abscisic acid, Festuca arundinacea, Eragrostis curvula, Sporobolus stapfianus     

Characterization of a wilty sunflower (Helianthus annuus L.) mutant II. Water relations, stomatal conductance, abscisic acid content in leaves and xylem sap of plants subjected to water deficiency

The response of w-1, a wilty sunflower (Helianthus annuus L.)mutant, to water stress is described in comparison with thecontrol line (W-1). Detached leaves of w-1 strongly dehydratedduring the first 30 min without significant changes in leafconductance, whereas W-1 responded rapidly to water loss byreducing stomatal aperture. After 2 h stress ABA increased slightlyin w-1, while W-1 leaves showed a 20-fold increase. When waterstress was imposed to potted plants by water withholding, w-1quickly dehydrated, and lost turgor, while W-1 maintained positiveturgor values for a longer period. Wild-type plants respondedto small changes in leaf water potential by accumulating ABAand by closing stomata, whereas in the mutant significant changesin ABA content and in stomatal conductance were found only atvery low water potentials. In another experiment in which waterwas withheld under high relative humidity, when soil water contentstarted to decrease W-1 rapidly closed stomata in the absenceof any change in leaf water status and the reduction in conductancewas paralleled by a rise in xylem sap ABA concentration. Bycontrast the mutant started to accumulate ABA in the xylem sapand to close stomata when soil water content and leaf waterpotential were dramatically reduced. The low endogenous ABAlevels and the inability to synthesize the hormone rapidly eitherin the leaves or in the roots seem to be responsible for thehigh sensitivity of w-1 to water stress. Key words: ABA, Helianthus annuus L, water relations, stomatal conductance, drought, wilty mutant     

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     

Effects of Periodical Soil Drying and Leaf Water Potential on the Sensitivity of Stomatal Response to Xylem ABA

The study on the changes of stomatal sensitivity in relation to xylem ABA during periodical soil drying and the effect of leaf water status on the stomatal sensitivity has confirmed that xylem ABA concentration is a good indicator of soil water status around roots and the relation between xylem ABA concentration and predawn leaf water potential remained constant during the three consecutive soil drying cycles based on the slopes of the fitted lines. The sensitivity of stomata to xylem ABA increased substantially as the soil drying cycles progressed, and the xylem ABA concentration needed to cause a 50% decrease of stomatal conductance was as low as 550 mnoL/L in the next two soil drying cycle, as compared with the 750 nmol/L ABA in the first cycle of soil drying. The results using the split-root system showed that leaf water deficit significantly enhanced the stomatal response to xylem ABA and the xylem ABA concentration needed to cause a 50% decrease in stomatal conductance was 2 to 4 times smaller in the whole-root-drying treatment than those in the semi-root- drying treatment. These results suggested that the sensitivity of stomata to xylem ABA concentration is not a fixed characteristic.     

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     

Can early wilting of old leaves account for much of the ABA accumulation in flooded pea plants?

When pea plants (Pisum sativum L.) were subjected to flooding,abscisic acid (ABA) content in shoots and roots increased upto 8-fold in the following days and stomatal conductance significantlydecreased. Although young leaves of flooded plants had a slightlyhigher water potential than those of the unflooded plants, oldleaves had lower water potential and lost turgor at the timewhen a substantial ABA increase was detected. In plants wherethe old leaves were clipped off, flooding did not cause anyABA increase during 7 d of the experimental period, except underconditions of higher transpiration demand, when the increasein ABA content was both delayed and small in scale (only I-fold).When intact plants were flooded and ABA was assayed separatelyin both old and young leaves, the ABA increase in old leavespreceded that in young leaves. Evidence here suggests that theflooding-induced ABA increase mainly results from the wiltingof old leaves. This suggests that young leaves may be protectedfrom wilting by ABA originating in old leaves under unfavourableenvironmental conditions. Key words: Waterlogging, soil flooding, ABA, leaf water relations, pea, Pisum sativum     

Abscisic acid produced in dehydrating roots may enable the plant to measure the water status of the soil

Abstract. Maize plants were grown in 1-m-long tubes of John Innes No. 2 potting compost. From the start of the experimental period, half of the plants were unwatered. Stomatal conductance of these plants was restricted 6 d after last watering and continued to decline thereafter. This was despite the fact that as a result of solute accumulation, unwatered plants showed consistently higher leaf turgors than well-watered plants. Leaf water potentials of unwatered plants were not significantly lower than those of plants that were watered well. Main seminal and nodal roots showed solute regulation in drying soil and continued to grow even in the driest soil, and plants growing in drying soil showed consistently higher root dry weights than did well-watered plants, water potentials and turgors of the tips of fine roots in the upper part of the column decreased as the soil dried. Soil drying below a water content of around 0–25 g cm⁻³ (a bulk soil water potential of between -0.2 and -0.3 MPa) resulted in a substantial increase in the ABA content of roots. As soil columns dried progressively from the top, ABA content increased in roots deeper and deeper in the soil. These responses suggest that ABA produced by dehydrating roots and which was subsequently transported to the shoots provided a sensitive indication of the degree of soil drying.     

Stomatal conductance in relation to xylem sap abscisic acid concentrations in two tropical trees, Acacia confusa and Litsea glutinosa

Two tropical trees, Acacia confusa and Litsea glutinosa, were grown under controlled conditions with their roots subjected to soil drying and soil compaction treatments. In both species, a decline in stomatal conductance resulting from soil drying took place much earlier than the decline of leaf water potential. Soil compaction treatment also resulted in a substantial decrease in stomatal conductance but had little effect on leaf water potential. A rapid and substantial increase in xylem abscisic acid (ABA) concenation ([ABA]), rather than hulk leaf ABA, was closely related to soil drying and soil compaction. A significant relationship between stomatal conductance (g_s) and xylem [ABA] was observed in both species. Artificially feeding ABA solutions to excised leaves of both species showed that the relationship bet ween g_s and [ABA] was very similar to that obtained from the whole plant, i.e. the relationship between g_s and xylem [ABA]. These results suggest that xylem ABA may act as a stress signal in the control of stomatal conductance.     

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.     

Root-to-shoot signalling when soil moisture is heterogeneous: increasing the proportion of root biomass in drying soil inhibits leaf growth and increases leaf abscisic acid concentration

To determine whether root-to-shoot signalling of soil moisture heterogeneity depended on root distribution, wild-type (WT) and abscisic acid (ABA)-deficient (Az34) barley (Hordeum vulgare) plants were grown in split pots into which different numbers of seminal roots were inserted. After establishment, all plants received the same irrigation volumes, with one pot watered (w) and the other allowed to dry the soil (d), imposing three treatments (1 d: 3 w, 2 d: 2 w, 3 d: 1 w) that differed in the number of seminal roots exposed to drying soil. Root distribution did not affect leaf water relations and had no sustained effect on plant evapotranspiration (ET). In both genotypes, leaf elongation was less and leaf ABA concentrations were higher in plants with more roots in drying soil, with leaf ABA concentrations and water potentials 30% and 0.2 MPa higher, respectively, in WT plants. Whole-pot soil drying increased xylem ABA concentrations, but maximum values obtained when leaf growth had virtually ceased (100 nm in Az34, 330 nm in WT) had minimal effects (<40% leaf growth inhibition) when xylem supplied to detached shoots. Although ABA may not regulate leaf growth in vivo, genetic variation in foliar ABA concentration in the field may indicate different root distributions between upper (drier) and lower (wetter) soil layers.     

Does ABA in the Xylem Control the Rate of Leaf Growth in Soil-Dried Maize and Sunflower Plants?

Maize (Zea mays L.) and sunflower (Helianthus annuus L.) plantswere grown in large volumes of soil and leaf growth rate wasmonitored on a daily basis. Half the plants were given a soildrying treatment and when they showed a significant restrictionof growth rate (compared to both their daily growth rate beforedrying and the average growth rate of well-watered plants onthe same day), leaf water relations were measured and xylemsap was extracted using several techniques. There was a significant negative log-linear relationship betweenthe rate of leaf growth and the concentration of ABA in thexylem for both species. There was no clear relationship betweenleaf growth rate and leaf water potential or turgor for eitherspecies. Assessment of different methods for sampling xylemsap suggests that exudates collected from stem stumps or samplescollected by pressurizing the whole root system are suitablefor estimating ABA concentration in xylem, at least with largeplants of maize or sunflower, provided the first few hundredcubic millimetres of collected sap are used for the assay. Centrifugationof sections of stems resulted in dilution of ABA in the xylemsap with sap squeezed from parenchyma tissue. This is because,at least in plants subjected to mild soil drying, the concentrationof the ABA in the xylem is far higher than that in the cellsap of stem tissue. Results support the proposal that ABA plays a major role asa chemical signal involved in the root-to-shoot communicationof the effects of soil drying. The non-hydraulic restrictionof leaf growth by a chemical signal can be explained by theextra root-sourced ABA in the xylem and may be an importantcomponent of the modification of growth and development whichresults from prolonged soil drought. Key words: Soil drying, ABA, leaf growth, Zea mays L., Helianthus annuus L.     

Are Roots a Source of Abscisic Acid for the Shoots of Flooded Pea Plants?

Flooding the soil for 2–5 d decreased stomatal conductancesof pea plants (Pisum sativum L., cv. Sprite) with six or sevenleaves. This coincided with slower transpiration, increasedleaf water potentials and increased concentrations of abscisicacid (ABA) in the leaves. No increase in ABA was found in theterminal 20 mm of roots of flooded plants over the same timeperiod. Small stomatal conductances associated with increases in foliarABA were also found in plants grown in nutrient solution whenaeration was halted, causing the equilibrium partial pressuresof dissolved oxygen to fall below 05 It Pa. No increase in ABAconcentration in young secondary roots of the non-aerated plantswas detected after 24, 48 or 72 h, even when the shoot, thepresumed site of deposition for any ABA from the roots, wasremoved 5–6 h before analysis. Similarly, ABA concentrations in roots were not increased whenthe nutrient solution was de-oxygenated by continuous purgingwith nitrogen gas. The abscisic acid concentration in leaf epidermis,the tissue most likely to be the recipient of any ABA movingin the transpiration stream from oxygen-deficient roots, waslower than in the remaining parts of the leaf when examinedin the mutant Argenteum which possesses easily removable epidermallayers. It is concluded that the leaves of plants subjectedto flooding of the soil or oxygen shortage in the root environmentare not enriched substantially with ABA from the roots. A moreprobable source of this growth regulator is the leaf itself. Key words: Pisum sativum, flooding, roots, hormones, aeration stress, abscisic acid, Argenteum mutant     

Characterization of a wilty sunflower (Helianthus annuus L.) mutant: III. Phenotypic interaction in reciprocal grafts from wilty mutant and wild-type plants

The present study was conducted to evaluate phenotypic interactionin reciprocal grafts between wilty (w-1) sunflower mutant andnormal (W-1) plants. The w-1 genotype is a ‘leaky’ABA-deficient mutant, characterized by high stomatal conductance,in both light and dark conditions, and high transpiration rate. In well-watered conditions, mutant scions grafted on to normalrootstock (w-1/W-1) showed higher leaf relative water content,leaf water potential and ABA levels than those of control grafts(w-1/w-1). In addition, detached leaves of w-1/W-1 exhibitedlower water loss than w-1/w-1 grafts, while mutant rootstockdid not affect the transpiration rate of detached W-1 leaves.When drought stress was imposed to potted plants by withholdingwater, the mutant scions grafted on to normal roots showed apartial phenotypic reversion. A rapid stomatal closure and arise in ABA levels in response to a small decrease in leaf waterpotential was observed. By contrast, in w-1/w-1 grafts significantreductions in stomatal conductance and ABA accumulation weredetected only in conjunction with a severe water deficit. W-1scions on mutant stocks (W-1/w-1) maintained the normal phenotypeof control wild-type grafts (W1/W-1). Key words: ABA, grafting, Helianthus annuus, stomatal conductance, water relations, wilty mutant     

Increased Synthesis of ABA in Partially Dehydrated Root Tips and ABA Transport from Roots to Leaves

Zhang, J. and Davies, W. J. 1987. Increased synthesis of ABAin partially dehydrated root tips and ABA transport from rootsto leaves.—J. exp. Bot. 38: 2015–2023. Isolated root tips of pea (Pisum sativum L. cv. Feltham First)and Commelina communis L. were air-dried until they lost between10% and 40% of their fresh weight, followed by a period of incubationat these reduced water contents. These treatments resulted inincreased ABA production, suggesting that root tips of bothspecies have the capacity to synthesize ABA in increased amountswhen water deficits develop in the root. The ABA concentrationin pea roots increased linearly as turgors fell below about0·15 M Pa and relative water contents (R WC) fell below90%. Commelina roots produced more ABA when RWC fell below asimilar value but the threshold turgor for increased ABA productionin Commelina roots was around 0·30 MPa. Roots of intact plants loaded with ABA as a result of incubationin solutions of varying concentrations provided ABA to leaveswhich resulted in increased ABA concentrations in the leaveswhen these were assayed several hours later. This occurred whenthese roots were not contributing substantially to transpirationalflux. Leaves on shoots that were enclosed and darkened and thereforenot transpiring, did not accumulate ABA from ‘loaded’roots. A role for root-sourced ABA in root-to-shoot communication ofthe effects of soil drying is discussed. Key words: ABA, roots, water relations     

Drought-induced Closure of Phaseolus vulgaris L. Stomata Precedes Leaf Water Deficit and any Increase in Xylem ABA Concentration

Young seedlings of two cultivars of Phaseolus vulgaris L. (cv.Cacahuate-72 and Michoacan-12A3) were subjected to a soil dryingtreatment. Stomata started to close before any leaf water deficitcould be detected. Soil drying promoted a small degree of ABAaccumulation in roots, but, at the time stomatal closure wasinitiated, at least in one cultivar, xylem ABA and bulk leafABA concentration were not enhanced. The mechanism of the stomatalresponse is discussed in terms of an as-yet unidentified regulatorof stomatal behaviour, redistribution of existing ABA in plants,and a high sensitivity of stomata to small changes in ABA concentration.     

Shoot induction of ABA-requiring genes in response to soil drying

Plant responses to water deficit are dynamic and varied, requiringco-ordination between the shoot and root. Among these responsesare alterations in gene expression. The expression of four genes,le4, le16, le20, and le25, which require increased ABA contentfor expression, was studied in tomato plants in which the rootsystems were divided between two large pots to impose waterdeficit gradually and to control signals from the root in responseto soil drying without inducing a signal from the shoot. Onegroup of plants had one-half of the roots watered, another grouphad both halves watered, and another group had neither halveswatered. In unwatered plants, the expression of le4 and le25correlated with ABA content, and that of le16 and le20 occurredbefore a detectable increase in leaf ABA content. The contrastingpatterns of expression indicate a difference in sensitivityof these genes to ABA or an additional signalling mechanism.Ample evidence indicates that shoot processes such as stomatalclosure are controlled by signals from the root. This studydemonstrates that genes may also be induced in the shoot bysignals from the root. Shoots of plants in which only half ofthe roots were watered showed no decrease in relative watercontent and no increase in ABA content; however, three of thefour genes, le4, le16, and le20, were induced. Root-to-shootcommunication plays a role in changes in gene expression andin alterations in physiological processes. Key words: Abscisic acid, water deficit, gene expression, split-root plants, long-distance signal