Decoupling the influence of leaf and root hydraulic conductances on stomatal conductance and its sensitivity to vapour pressure deficit as soil dries in a drained loblolly pine plantation

The study examined the relationships between whole tree hydraulic conductance ( K _tree) and the conductance in roots ( K _root) and leaves ( K _leaf) in loblolly pine trees. In addition, the role of seasonal variations in K _root and K _leaf in mediating stomatal control of transpiration and its response to vapour pressure deficit ( D ) as soil-dried was studied. Compared to trunk and branches, roots and leaves had the highest loss of conductivity and contributed to more than 75% of the total tree hydraulic resistance. Drought altered the partitioning of the resistance between roots and leaves. As soil moisture dropped below 50%, relative extractable water (REW), K _root declined faster than K _leaf. Although K _tree depended on soil moisture, its dynamics was tempered by the elongation of current-year needles that significantly increased K _leaf when REW was below 50%. After accounting for the effect of D on g _s, the seasonal decline in K _tree caused a 35% decrease in g _s and in its sensitivity to D , responses that were mainly driven by K _leaf under high REW and by K _root under low REW. We conclude that not only water stress but also leaf phenology affects the coordination between K _tree and g _s and the acclimation of trees to changing environmental conditions.     

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.     

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.     

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.     

Elevated CO2 concentrations and stomatal density: observations from 17 plant species growing in a CO2 spring in central Italy

Stomatal density (SD) and stomatal conductance ( g _s) can be affected by an increase of atmospheric CO₂ concentration. This study was conducted on 17 species growing in a naturally enriched CO₂ spring and belonging to three plant communities. Stomatal conductance, stomatal density and stomatal index (SI) of plants from the spring, which were assumed to have been exposed for generations to elevated [CO₂], and of plants of the same species collected in a nearby control site, were compared. Stomatal conductance was significantly lower in most of the species collected in the CO₂ spring and this indicated that CO₂ effects on g _s are not of a transitory nature but persist in the long term and through plant generations. Such a decrease was, however, not associated with changes in the anatomy of leaves: SD was unaffected in the majority of species (the decrease was only significant in three out of the 17 species examined), and also SI values did not vary between the two sites with the exception of two species that showed increased SI in plants grown in the CO₂-enriched area. These results did not support the hypothesis that long-term exposure to elevated [CO₂] may cause adaptive modification in stomatal number and in their distribution.     

Decreased root hydraulic conductivity reduces leaf water potential, initiates stomatal closure and slows leaf expansion in flooded plants of castor oil (Ricinus communis) despite diminished delivery of ABA from the roots to shoots in xylem sap

Soil flooding reduced stomatal conductance (gs) and slowed transpiration, CO₂ uptake and leaf elongation in Ricinus communis within 2–6 h. These flood-induced responses developed further over the next 21 h. They were not associated with increased delivery of abscisic acid (ABA) in xylem sap. Instead, ABA delivery from flooded roots decreased 6-fold within 3 h, and remained low thereafter. Root hydraulic conductance (Lp) was depressed 47% below control values within 2 h of soil flooding, and declined further during the next 21 h. The smaller Lp temporarily decreased leaf water potentials (ΨL) by up to −0.4 MPa, and caused visible wilting 3 h into the flooding treatment at 80% relative humidity. Consequently, ABA concentrations in the shoot were increased, as indicated by analyses of phloem sap. Wilting, fall in ΨL and a reduction in gs were delayed for 6 h when 0.6 MPa pneumatic pressure (technical maximum) was applied to the roots. In flooded plants, phloem sap ABA concentrations returned to normal after 24 h. The initial stomatal closure, caused by soil flooding in R. communis , is attributed to decreased leaf hydration arising from the reduced LP of oxygen-deficient roots. Continued stomatal closure and slow leaf expansion beyond 24 h were presumably achieved by non-hydraulic means.     

Sap salinity effects on xylem conductivity in two mangrove species

Xylem sap salinity and conductivity were examined in two mangrove ecosystem tree species . For Avicennia germinans , extracted xylem sap osmotic potentials ranged from −0.24 to −1.36 MPa versus −0.14 to −0.56 MPa for Conocarpus erectus. Xylem sap of Conocarpus did not vary in osmotic potential between sites nor between predawn and midday. In Avicennia , values were more negative at midday than predawn, and also more negative at hypersaline than hyposaline sites. After removing embolisms, specific conductivity ( K _s) was measured as a function of salinity of the artificial xylem sap perfusion. For both species the lowest K _s values, about 70% of the maximum K _s, were obtained when stems were perfused with deionized water (0 m m ; 0.0 MPa) or with a 557-m m saline solution (−2.4 MPa). Higher K _s values were obtained in the range from −0.3 to −1.2 MPa, with a peak at −0.82 ± 0.08 MPa for Avicennia and −0.75 ± 0.08 MPa for Conocarpus . The variations in K _s values with minima both at very low and very high salt concentrations were consistent with published results for swelling and shrinking of synthetic hydrogels, suggesting native hydrogels in pit membranes of vessels could help regulate conductivity.     

Relationships between sap flow and water potential in woody or perennial plants on islands of the Great Barrier Reef

Abstract. This paper describes studies on trees of Pisonia grandis , bushes of Argusia argentea , and the perennial herb Melanthera biflora , growing on One Tree Island, a coral cay of the Great Barrier Reef with 'soil' of coarse coral rubble. Water potential (Ψ_b, measured on small shoots with a pressure chamber), sap flow, stomatal conductance, vapour pressure deficit and photon flux density were monitored over day/night cycles. Sap flow and Ψ_b responded to changes in light and humidity. From these experiments good linear correlations were found between sap flow in a shoot and Ψ_b of similar adjacent shoots. The linearity suggests that the resistance to sap flow is constant as Ψ_b varies. The correlation, however, does not indicate a causal relationship between Ψ_b of an individual shoot on the plant and its sap flow. Ψ_b was only slightly different in shaded shoots from those in sunshine, although sap flow would be expected to differ between them. Enclosing shoots and so reducing their transpiration and sap flow to very low rates resulted in only small changes in Ψ_b of the enclosed shoots; Tb of such enclosed shoots should closely approximate that of the xylem at the point of shoot attachment. From these results it is suggested that the resistance to water flow in shoot and leaf xylem is small compared to the resistance further down the plant, in the root or at the root/soil interface. Shoot xylem water potential would be similar for all parts of the plant, and in such plants the water potential of shoots in the shade would be determined by the overall water use of the plant.     

Fluoride inhibits root water transport and affects leaf expansion and gas exchange in aspen (Populus tremuloides) seedlings

The effects of sodium fluoride (0.3, 5 and 10 m M NaF) on root hydraulic conductivity, and gas exchange processes were examined in aspen ( Populus tremuloides Michx.) seedlings grown in solution culture. A long-term exposure of roots to NaF significantly decreased root hydraulic conductivity ( L _p) and stomatal conductance ( g _s). Root absorbed NaF significantly affected electrolyte leakage in leaf tissues and substantially restricted leaf expansion. NaF did not significantly affect leaf chlorophyll contents but decreased net photosynthesis ( P _n). A short-term exposure of excised roots to 5 m M NaF and KF significantly decreased root water flow ( Q _v) with a concomitant decline in root respiration and reduced g _s when applied through intact roots or excised stems. The same molar concentration of NaCl also decreased Q _v and g _s in intact seedlings, but to a lesser extent than NaF or KF, and did not significantly affect root respiration. The results suggest that fluoride metabolically inhibited Q _v or L _p, probably by affecting water channel activity. We suggest that the metabolic inhibition of L _p by root-absorbed fluoride affected gas exchange and leaf expansion in aspen seedlings.     

Responses of apple leaf stomata: a model for single leaves and a whole tree

Abstract. An empirical model of stomatal response to environmental factors was developed from measurements of stomatal conductance ( g _s) made in a leaf chamber under controlled conditions. Results presented in a companion paper (Warrit, Landsberg & Thorpe, 1980) indicated that the model could be written in terms of only two factors, photon flux density ( Q _p) and leaf to air vapour pressure gradient ( D ). The response of Q _p was hyperbolic and that to D linear; combining these the equation of the model is where g _r is a reference conductance, α is the slope of the response to D and β indicates the sensitivity of g _s response to Q _p. Values of α were 0.20 and 0.30 kPa⁻¹ in June and August; the corresponding values of β were 59 and 79 μmol m⁻² s⁻¹.
The model was tested against mean values of g _s obtained with a porometer in the field, using environmental measurements as inputs. Correspondence between measured and calculated values was good. Transpiration rates were calculated from the Penman-Monteith equation, with stomatal resistance values calculated from the model, and compared with gravimetric measurements of tree water use. It was shown that transpiration could be calculated with acceptable accuracy. The effects of variations in stomatal resistance on transpiration rates under a range of conditions were explored using the model and the Penman- Monteith equation.     

Chemical regulation of gas exchange and growth of plants in drying soil in the field

There is now substantial evidence that chemical regulation ofshoot physiology occurs in droughted plants in the field. Theevidence that ABA may play a role in such regulation is considered,and topics of relevance to the worker interested in determiningthe ABA relations of plants in the field; such as the methodsused for ABA quantification, the relevance of quantifying ABAin various plant tissues, methods of xylem sap collection andtiming of sap collection are reviewed. A possible role of tissuesensitivity to ABA in controlling the diurnal changes in stomatalconductance and leaf growth rate seen in the field is also considered. Key words: ABA, drought, stomatal conductance, leaf growth, hormonal sensitivity, xylem sap     

An external heat pulse method for measurement of sap flow through fruit pedicels, leaf petioles and other small-diameter stems

The external heat ratio method is described for measurement of low rates of sap flow in both directions through stems and other plant organs, including fruit pedicels, with diameters up to 5 mm and flows less than 2 g h⁻¹. Calibration was empirical, with heat pulse velocity ( v _h) compared to gravimetric measurements of sap flow. In the four stem types tested ( Actinidia sp. fruit pedicels, Schefflera arboricola petioles, Pittosporum crassifolium stems and Fagus sylvatica stems), v _h was linearly correlated with sap velocity ( v _s) up to a v _s of approximately 0.007 cm s⁻¹, equivalent to a flow of 1.8 g h⁻¹ through a 3-mm-diameter stem. Minimum detectable v _s was approximately 0.0001 cm s⁻¹, equivalent to 0.025 g h⁻¹ through a 3-mm-diameter stem. Sensitivity increased with bark removal. Girdling had no effect on short-term measurements of in vivo sap flow, suggesting that phloem flows were too low to be separated from xylem flows. Fluctuating ambient temperatures increased variability in outdoor sap flow measurements. However, a consistent diurnal time-course of fruit pedicel sap flow was obtained, with flows towards 75-day-old kiwifruit lagging behind evaporative demand and peaking at 0.3 g h⁻¹ in the late afternoon.     

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

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

The effect of competition from neighbours on stomatal conductance in lettuce and tomato plants

Competition decreased transpiration from young lettuce plants after 2 days, before any reductions in leaf area became apparent, and stomatal conductance (g(s) ) of lettuce and tomato plants was also reduced. Stomatal closure was not due to hydraulic signals or competition for nutrients, as soil water content, leaf water status and leaf nitrate concentrations were unaffected by neighbours. Competition-induced stomatal closure was absent in an abscisic acid (ABA)-deficient tomato mutant, flacca, indicating a fundamental involvement of ABA. Although tomato xylem sap ABA concentrations were unaffected by the presence of neighbours, ABA/pH-based stomatal modulation is still likely to underlie the response to competition, as soil and xylem sap alkalization was observed in competing plants. Competition also modulated leaf ethylene production, and treatment of lettuce plants with an ethylene perception inhibitor (1-methylcyclopropene) diminished the difference in g(s) between single and competing plants grown in a controlled environment room, but increased it in plants grown in the greenhouse: ethylene altered the extent of the stomatal response to competition. Effects of competition on g(s) are discussed in terms of the detection of the absence of neighbours: increases in g(s) and carbon fixation may allow faster initial space occupancy within an emerging community/crop.     

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

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

Keeping a positive carbon balance under adverse conditions: responses of photosynthesis and respiration to water stress

Drought and salinity (i.e. soil water stress) are the main environmental factors limiting photosynthesis and respiration and, consequently, plant growth. This review summarizes the current status of knowledge on photosynthesis and respiration under water stress. It is shown that diffusion limitations to photosynthesis under most water stress conditions are predominant, involving decreased mesophyll conductance to CO₂, an important but often neglected process. A general failure of photochemistry and biochemistry, by contrast, can occur only when daily maximum stomatal conductance ( g _s) drops below 0.05–0.10 mol H₂O m⁻² s⁻¹. Because these changes are preceded by increased leaf antioxidant activities ( g _s below 0.15–0.20 mol H₂O m⁻² s⁻¹), it is suggested that metabolic responses to severe drought occur indirectly as a consequence of oxidative stress, rather than as a direct response to water shortage. As for respiration, it is remarkable that the electron partitioning towards the alternative respiration pathway sharply increases at the same g _s threshold, although total respiration rates are less affected. Despite the considerable improvement in the understanding of plant responses to drought, several gaps of knowledge are highlighted which should become research priorities for the near future. These include how respiration and photosynthesis interact at severe stress, what are the boundaries and mechanisms of photosynthetic acclimation to water stress and what are the factors leading to different rates of recovery after a stress period.     

Enhanced assimilation rate and water use efficiency with latitude through increased photosynthetic capacity and internal conductance in balsam poplar (Populus balsamifera L.)

In outdoor common gardens, high latitude populations of deciduous tree species often display higher assimilation rates ( A ) than low latitude populations, but they accomplish less height. To test whether trends in A reflect adaptation to growing season length or, alternatively, are garden growth artefacts, we examined variation in height increment and ecophysiological traits in a range-wide collection of Populus balsamifera L. populations from 21 provenances, during unconstrained growth in a greenhouse. Rooted cuttings, maintained without resource limitation under 21 h photoperiod for 90 d, displayed increasing height growth, A , leaf mass per area and leaf N per area with latitude whereas stomatal conductance ( g _s) showed no pattern. Water-use efficiency as indicated by both gas exchange and δ ¹³C increased with latitude, whereas photosynthetic nitrogen-use efficiency decreased. Differences in δ ¹³C were less than expected based on A/g _s, suggesting coextensive variation in internal conductance ( g _m). Analysis of A – C _i curves on a subset of populations showed that high latitude genotypes had greater g _m than low-latitude genotypes. We conclude that higher peak rates of height growth in high latitude genotypes of balsam poplar are supported by higher A , achieved partly through higher g _m, to help compensate for a shorter growing season.     

Linking physiological processes with mangrove forest structure: phosphorus deficiency limits canopy development, hydraulic conductivity and photosynthetic carbon gain in dwarf Rhizophora mangle

Spatial gradients in mangrove tree height in barrier islands of Belize are associated with nutrient deficiency and sustained flooding in the absence of a salinity gradient. While nutrient deficiency is likely to affect many parameters, here we show that addition of phosphorus (P) to dwarf mangroves stimulated increases in diameters of xylem vessels, area of conductive xylem tissue and leaf area index (LAI) of the canopy. These changes in structure were consistent with related changes in function, as addition of P also increased hydraulic conductivity ( K _s), stomatal conductance and photosynthetic assimilation rates to the same levels measured in taller trees fringing the seaward margin of the mangrove. Increased xylem vessel size and corresponding enhancements in stem hydraulic conductivity in P fertilized dwarf trees came at the cost of enhanced midday loss of hydraulic conductivity and was associated with decreased assimilation rates in the afternoon. Analysis of trait plasticity identifies hydraulic properties of trees as more plastic than those of leaf structural and physiological characteristics, implying that hydraulic properties are key in controlling growth in mangroves. Alleviation of P deficiency, which released trees from hydraulic limitations, reduced the structural and functional distinctions between dwarf and taller fringing tree forms of Rhizophora mangle .     

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.