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.     

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.     

Stomatal control by both [ABA] in the xylem sap and leaf water status: a test of a model for draughted or ABA-fed field-grown maize

A model of maize stomatal behaviour has been developed, in which stomatal conductance is linked to the concentration of abscisic acid ([ABA]) in the xylem sap, with a sensitivity dependent upon the leaf water potential (Ψ₁). It was tested against two alternative hypotheses, namely that stomatal sensitivity to xylem [ABA] would be linked to the leaf-to-air vapour pressure difference (VPD), or to the flux of ABA into the leaf. Stomatal conductance (g_s) was studied: (1) in field-grown plants whose xylem [ABA] and Ψ₁ depended on soil water status and evaporative demand; (2) in field-grown plants fed with ABA solutions such that xylem [ABA] was artificially raised, thereby decreasing g_s and increasing Ψ₁ and leaf-to-air VPD; and (3) in ABA-fed detached leaves exposed to varying evaporative demands, but with a constant and high Ψ₁. The same relationships between g_s, xylem [ABA] and Ψ₁, showing lower stomatal sensitivity to [ABA] at high Ψ₁, applied whether variations in xylem [ABA] were due to natural increase or to feeding, and whether variations in Ψ₁, were due to changes in evaporative demand or to the increased Ψ₁ observed in ABA-fed plants. Conversely, neither the leaf-to-air VPD nor the ABA flux into the leaf accounted for the observed changes in stomatal sensitivity to xylem [ABA]. The model, using parameters calculated from previous field data and the detached-leaf data, was tested against the observations of both ABA-fed and droughted plants in the field. It accounted with reasonable accuracy for changes in g_s (r² ranging from 0.77 to 0.81). These results support the view that modelling of stomatal behaviour requires consideration of both chemical and hydraulic aspects of root-to-shoot communication.     

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.     

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.     

ABA xylem concentrations determine maximum daily leaf conductance of field-grown Vitis vinifera L. plants

Differences in maximum leaf conductance in grapevine plants growing in soils with contrasting water availabilities during mid-summer in Portugal could be accounted for by differences in the concentration of ABA in xylem sap. This conclusion is reinforced by the observation that the relationship between leaf conductance and endogenous ABA concentration can be mimicked by the application of exogenous ABA to leaves detached from irrigated plants. During the day, leaf conductance decreased after a morning peak, even when the leaves remained in a constant environment at a moderate temperature and leaf-to-air vapour pressure difference. This decline in leaf conductance was not a consequence of an increase in the xylem ABA concentration or the rate of delivery of this compound by the transpiratory stream. The afternoon depression in leaf conductance was associated with an apparent limitation in stomatal opening potential, which persisted even when detached leaves were fed with water and rehydrated. The reason for this inhibition has still to be identified.     

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.     

Integration of hydraulic and chemical signalling in the control of stomatal conductance and water status of droughted plants

We describe here an integration of hydraulic and chemical signals which control stomatal conductance of plants in drying soil, and suggest that such a system is more likely than control based on chemical signals or water relations alone. The determination of xylem [ABA] and the stomatal response to xylem [ABA] are likely to involve the water flux through the plant. (1) If, as seems likely, the production of a chemical message depends on the root water status (Ψ_r), it will not depend solely on the soil water potential (Ψ_s) but also on the flux of water through the soil-plant-atmosphere continuum, to which are linked the difference between Ψ_r and Ψ_s. (2) The water flux will also dilute the concentration of the message in the xylem sap. (3) The stomatal sensitivity to the message is increased as leaf water potential falls. Stomatal conductance, which controls the water flux, therefore would be controlled by a water-flux-dependent message, with a water-flux-dependent sensitivity. In such a system, we have to consider a common regulation for stomatal conductance, leaf and root water potentials, water flux and concentration of ABA in the xylem. In order to test this possibility, we have combined equations which describe the generation and effects of chemical signals and classical equations of water flux. When the simulation was run for a variety of conditions, the solution suggested that such common regulation can operate. Simulations suggest that, as well as providing control of stomatal conductance, integration of chemical and hydraulic signalling may also provide a control of leaf water potential and of xylem [ABA], features which are apparent from our experimental data. We conclude that the root message would provide the plant with a means to sense the conditions of water extraction (soil water status and resisance to water flux) on a daily timescale, while the short-term plant response to this message would depend on the evaporative demand.     

What information is conveyed by an ABA signal from maize roots in drying field soil?

During two seasons, ABA concentrations were monitored in roots, leaves and xylem sap of field-grown maize. The water status of soil and plant was also measured. Plants were grown on plots with compacted or non-compacted soil, which were irrigated or remained unwatered. ABA concentration in the xylem sap before dawn and in the roots increases 25-fold and five-fold, respectively, as the soil dried, with a close correlation with the soil water status, but with no clear effect of the soil structure. In contrast to the results of several laboratory experiments, no appreciable increase in xylem [ABA] and reduction in stomatal conductance were observed with dehydration of the part of the root system located in soil upper layers. These responses only occurred when the water reserve of the whole soil profile was close to depletion and the transpiration declined. Xylem [ABA] measured during the day was appreciably higher in the compacted treatment than in non-compacted treatment, unlike that measured before dawn. Since a mechanical message is unlikely to undergo such day-night alterations, we suggest that this was due to a faster decrease in root water potential and water flux in the compacted treatment, linked to the root spatial arrangement. These results raise the possibility that ABA concentration in the xylem sap could be controlled by two coexisting mechanisms: (1) the rate of ABA synthesis in the roots linked to the soil or root water status, as shown in laboratory experiments; (2) the dilution of ABA in the water flow from roots, which could be an overriding mechanism in field conditions. This second mechanism would allow the plant to sense the water flux through the root system.     

Stomatal control by fed or endogenous xylem ABA in sunflower: interpretation of correlations between leaf water potential and stomatal conductance in anisohydric species

The stomatal conductance of several anisohydric plant species, including field-grown sunflower, frequently correlates with leaf water potential (φ₁), suggesting that chemical messages travelling from roots to shoots may not play an important role in stomatal control. We have performed a series of experiments in which evaporative demand, soil water status and ABA origin (endogenous or artificial) were varied in order to analyse stomatal control. Sunflower plants were subjected to a range of soil water potentials under contrasting air vapour pressure deficits (VPD, from 0.5 to 2.5 kPa) in the field, in the glasshouse or in a humid chamber. Sunflower plants were also fed through the xylem with varying concentrations of artificial ABA, in the glasshouse and in the field. Finally, detached leaves were fed directly with varying concentrations of ABA under three contrasting VPDs. A unique relationship between stomatal conductance (g_s) and the concentration of ABA in the xylem sap (xylem [ABA]) was observed in all cases. In contrast, the relationship between φ₁ and g_s varied substantially among experiments. Its slope was positive for droughted plants and negative for ABA-fed whole plants or detached leaves, and also varied appreciably with air VPD. All observed relationships could be modelled on the basis of the assumption that φ₁ had no controlling effect on g_s. We conclude that stomatal control depended only on the concentration of ABA in the xylem sap, and that φ₁ was controlled by water flux through the plant (itself controlled by stomatal conductance). The possibility is also raised that differences in stomatal ‘strategy’ between isohydric plants (such as maize, where daytime φ₁ does not vary appreciably with soil water status) and anisohydric plants (such as sunflower) may be accounted for by the degree of influence of φ₁ on stomatal control, for a given level of xylem [ABA]. We propose that statistical relationships between φ₁ and g_s are only observed when φ₁ has no controlling action on stomatal behaviour.     

Drought adaptation of field grown wheat in relation to soil physical conditions

In order to investigate the effects of soil texture on possible non-hydraulic signals under field conditions, spring wheat plants (Triticum aestivum L. cv. Cadensa) grown in sand and loam soils and with a well developed root system were exposed to slow soil drying in the late vegetative stage of growth. Soil water potential and content were measured daily at different depths and plant responses were measured in flag leaves. When the average soil water potential in the top soil layers (0–25 cm depth in sand and 0–45 cm depth in loam) dropped to –60 or –70 kPa and the lower soil layers were still at field capacity, morning xylem [ABA] (0.03–0.04 vs. 0.06–0.08 mmol m-3) and midday leaf ABA concentration increased (250–300 vs. 400–450 ng/g DW) and leaf conductance decreased relatively to well-watered (control) plants (0.75–0.88 vs. 0.64–0.70 mol m-2 s-1). These responses took place before any decrease in leaf water potential occurred as compared with control plants, indicating that they were triggered by root-borne signals due to reduced root water status in the top soil layers. At this stage the soil water content was as low as 6% by volume, the fraction of roots in ‘wet’ soil was 0.12 and relative available soil water was 45% in sand and still high 20%, 0.48 and 70%, respectively, in loam of the whole soil profile indicating that roots were responding to soil water availability and not soil water content at a certain evaporative demand. In addition, similar responses occurred at high and low evaporative demands (3.4–5.2 vs. 0.6–4.0 mm/day of potential evapotranspiration). This revised version was published online in July 2006 with corrections to the Cover Date.     

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 stomatal closure caused by xylem ABA explain the inhibition of leaf photosynthesis under soil drying?

Effects of leaf water deficit and increase in endogenous ABA on photosynthesis of two tropical trees, t Acacia confusa and t Leucaena leucocephala, were investigated with two soil-drying methods, i.e. half or whole root system was subjected to soil drying. Half-root drying was achieved by allowing upper layer of soil column to dry and lower layer of soil column to remain watered. Half-root drying had little effect on leaf water potential, but when compared to the well-watered control, both methods of soil drying substantially increased the ABA concentration in xylem and reduced leaf conductance in both species. There was a significant relationship between leaf conductance and xylem ABA concentrations in both species, which was comparable to the same relationship that was generated by feeding ABA to excised twigs. The rate of photosynthesis was inhibited substantially in both soil-drying treatments and in both species, but photochchemical efficiency, measured as a ratio of variable fluorescence to a peak fluorescence emission of a dark-adapted leaf (F_v/F_m), was not reduced except in the whole root-dried t L. leucocephala plants where leaf water potential was reduced to –2.5 MPa. In all the cases where photosynthesis was inhibited, there was a concomitant reduction in both leaf conductance and calculated internal CO₂ concentration. After two days of rewatering, leaf water potential and xylem ABA concentration rapidly returned to pre-treatment levels, but leaf conductance and photosynthesis of both whole-root and half root dried t L. leucocephala remained inhibited substantially. Rewatering led to a full recovery of both stomatal conductance and photosynthesis in soil-dried t A. confusa, although its photosynthesis of whole-root dried plants did not recover fully but such difference was not significant statistically. These results suggest that drought-induced decline of photosynthesis was mainly a result of the stomatal factor caused by the increase of ABA concentration in the xylem sap. Non-stomatal factors, e.g. reduced photochemical activity and/or carbon metabolic activity, were species-specific and were brought about only at very low water potential.     

Accounting for sap flow from different parts of the root system improves the prediction of xylem ABA concentration in plants grown with heterogeneous soil moisture

When soil moisture is heterogeneous, sap flow from, and ABA status of, different parts of the root system impact on leaf xylem ABA concentration ([X-ABA]leaf). The robustness of a model for predicting [X-ABA]leaf was assessed. 'Two root-one shoot' grafted sunflower (Helianthus annuus L.) plants received either deficit irrigation (DI, each root system received the same irrigation volumes) or partial rootzone drying (PRD, only one root system was watered and the other dried the soil). Irrespective of whether relative sap flow was assessed using sap flow sensors in vivo or by pressurization of de-topped roots, each root system contributed similarly to total sap flow during DI, while sap flow from roots in drying soil declined linearly with soil water potential (Psisoil) during PRD. Although Psisoil of the irrigated pot determined the threshold Psisoil at which sap flow from roots in drying soil decreased, the slope of this decrease was independent of the wet pot Psisoil. Irrespective of whether sap was collected from the wet or dry root system of PRD plants, or a DI plant, root xylem ABA concentration increased as Psisoil declined. The model, which weighted ABA contributions of each root system according to the sap flow from each, almost perfectly explained [X-ABA] immediately above the graft union. That the model overestimated measured [X-ABA]leaf may result from changes in [X-ABA] along the transport pathway or an artefact of collecting xylem sap from detached leaves. The implications of declining sap flow through partially dry roots during PRD for the control of stomatal behaviour and irrigation scheduling are discussed.     

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     

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。     

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．     

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.     

Long-distance signals regulating stomatal conductance and leaf growth in tomato (Lycopersicon esculentum) plants subjected to partial root-zone drying

Tomato (Lycopersicon esculentum Mill. cv. Ailsa Craig) plants were grown with roots split between two soil columns. After plant establishment, water was applied daily to one (partial root-zone drying-PRD) or both (well-watered control-WW) columns. Water was withheld from the other column in the PRD treatment, to expose some roots to drying soil. Soil and plant water status were monitored daily and throughout diurnal courses. Over 8 d, there were no treatment differences in leaf water potential (psileaf) even though soil moisture content of the upper 6 cm (theta) of the dry column in the PRD treatment decreased by up to 70%. Stomatal conductance (gs) of PRD plants decreased (relative to WW plants) when of the dry column decreased by 45%. Such closure coincided with increased xylem sap pH and did not require increased xylem sap abscisic acid (ABA) concentration ([X-ABA]). Detached leaflet ethylene evolution of PRD plants increased when of the dry column decreased by 55%, concurrent with decreased leaf elongation. The physiological significance of enhanced ethylene evolution of PRD plants was examined using a transgenic tomato (ACO1AS) with low stress-induced ethylene production. In response to PRD, ACO1AS and wild-type plants showed similar xylem sap pH, [X-ABA] and gs, but ACO1AS plants showed neither enhanced ethylene evolution nor significant reductions in leaf elongation. Combined use of genetic technologies to reduce ethylene production and agronomic technologies to sustain water status (such as PRD) may sustain plant growth under conditions where yield would otherwise be significantly reduced.