Stomatal conductance,growth and root signaling in Betula pendula seedlings subjected to partial soil drying

Seedlings of Betula pendula Roth were grown with their root systems separated between two soil compartments. Four treatments were imposed: (i) adequate irrigation in both compartments (WW, controls); (ii) adequate irrigation in one compartment and drought in the other compartment (WD); (iii) drought in both compartments (DD); and (iv) half of the root system severed and the remainder kept well-watered (root excision, RE). Predawn leaf water potential, stomatal conductance, soil-to-leaf specific hydraulic conductance, and root and leaf growth decreased in DD-treated seedlings, which also displayed severe leaf shedding (30% loss in leaf area). The DD treatment also resulted in increased concentrations of abscisic acid (ABA) and its glucose ester in the xylem sap of roots and shoots compared to concentrations in control seedlings (about 200 versus 20 nM). Despite the difference in xylem sap concentrations, total ABA flux to the shoots was similar in the two treatments (1-2 pmol ABA m(-2) leaf area s(-1)) as a result of reduced transpiration in the DD-treated seedlings. Compared with root growth in control plants, root growth increased in the RE-treated plants and decreased in the drying compartment of the WD treatment; however, the RE and WD treatments only slightly reduced leaf expansion, and had no detectable effects on shoot water relations or ABA concentrations of the root and shoot xylem sap. We conclude that short-term soil water depletion affecting only 50% of the root system does not cause a measurable stress response in birch shoots, despite root growth cessation in the fraction of drying soil.     

Stomatal and leaf growth responses to partial drying of root tips in willow

Root tips of intact willow (Salix dasyclados Wimm., Clone 81-090) plants were partially dried by exposure to ambient greenhouse air and then kept in water-vapor-saturated air for up to 3 days. The drying treatment increased abscisic acid (ABA) concentrations in both the root tips subjected to drying and in the xylem sap, while it reduced leaf stomatal conductance and leaf extension rate. Despite the decrease in stomatal conductance, leaf water potentials were unaffected by the root drying treatment, indicating that the treatment reduced hydraulic conductivity between roots and foliage. After roots subjected to drying were returned to a nutrient solution or excised, ABA concentrations in the remaining roots and in the xylem sap, stomatal conductance of mature leaves and extension rate of unfolding leaves all returned to values observed in control plants. The 4-fold increase in xylem sap ABA concentration following the root drying treatment was not solely the result of reduced sap flow, and thus may be considered a potential cause, not merely a consequence, of the observed reduction in stomatal conductance.     

Modulation of leaf conductance by root to shoot signaling under water stress in Arabidopsis

Signal communication between root and shoot plays a crucial role in plant resistance to water stress. While many studies on root to shoot signals have been carried out in many plant species, no information is available for the model plant, Arabidopsis, whose adoption has great significance for further probing the molecular aspects of long distance stress signals. Here, we introduced the establishment of techniques for investigations of root to shoot signals in Arabidopsis. Stomatal movements in relation to root signals were probed by using these techniques. The results show that Arabidopsis is a suitable plant species for partial roots drying (PRD) experiments. In the PRD system, while no significant differences were found in leaf water potential between well-watered and stressed plants, water stress led to a decrease in leaf conductance, which suggests a regulation of stomatal movements by root to shoot signals. While water stress caused a significant increase in the concentration of sap abscisic acid (ABA) of xylem, no increase in xylem sap pH was observed. Moreover, the increase in the ABA content of xylem coincided with the decrease in leaf conductance, which suggests a possible role of ABA in the regulation of stomatal movements. Infrared temperature images showed that leaf temperatures of PRD plant were higher compared with those of well-watered plants, which further indicates that stomatal movements can be modulated by root signals. The confirmation of root to shoot signaling in Arabidopsis has established a basis for further investigation into the molecular mechanisms of the root to shoot signaling under water stress.     

Drought tolerance,xylem sap abscisic acid and stomatal conductance during soil drying: a comparison of canopy trees of three temperate deciduous angiosperms

Patterns of water relations, xylem sap abscisic acid concentration ([ABA]) and stomatal aperture were characterized and compared in drought-sensitive black walnut (Juglans nigra L.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.) trees co-occurring in a second-growth forest in Missouri, USA. There were strong correlations among reduction in predawn leaf water potential, increased xylem sap [ABA] and stomatal closure in all species. Stomatal conductance was more closely correlated with xylem sap ABA concentration than with ABA flux or xylem sap pH and cation concentrations. In isohydric black walnut, increased concentrations of ABA in the xylem sap appeared to be primarily of root origin, causing stomatal closure in response to soil drying. In anisohydric sugar maple and white oak, however, there were reductions in midday leaf water potential associated with stomatal closure, making it uncertain whether drought-induced xylem sap ABA was of leaf or root origin. The role of root-originated xylem sap ABA in these species as a signal to the shoot of the water status of the roots is, therefore, less certain.     

Drought tolerance,xylem sap abscisic acid and stomatal conductance during soil drying: a comparison of young plants of four temperate deciduous angiosperms

Patterns of water relations, xylem sap abscisic acid (ABA) concentration ([ABA]) and stomatal aperture were compared in drought-sensitive black walnut (Juglans nigra L.) and black willow (Salix nigra Marsh.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.). Strong correlations among reduction in predawn water potential, increase in xylem sap [ABA] and stomatal closure were observed in all species. Stomatal response was more highly correlated with xylem [ABA] than with ABA flux. Xylem sap pH and ion concentrations appeared not to play a major role in the stomatal response of these species. Stomata were more sensitive to relative changes in [ABA] in drought-sensitive black walnut and black willow than in sugar maple and white oak. In the early stages of drought, increased [ABA] in the xylem sap of black walnut and black willow was probably of root origin and provided a signal to the shoot of the water status of the roots. In sugar maple and white oak, leaf water potential declined with the onset of stomatal closure, so that stomatal closure also may have occurred in response to the change in leaf water potential.     

Canopy stomatal conductance and xylem sap abscisic acid (ABA) in mature Scots pine during a gradually imposed drought

We investigated the effect of drought on canopy stomatal conductance (g(c)), and examined the hypothesis that g(c) is controlled by the chemical messenger abscisic acid (ABA) produced in roots. Beginning in November 1994, we subjected a mature stand of Scots pine (Pinus sylvestris L.) to an imposed 11-month drought. Control plots were maintained at average-season soil water content. Xylem sap was extracted from shoots at regular intervals from April to November 1995. Soil water, sap flow and leaf water potentials (predawn to dusk) were recorded at the same time. Canopy stomatal conductance was calculated from sap flow data and xylem sap ABA concentrations ([ABA(xyl)]) were measured by radioimmunoassay. Mean [ABA(xyl)] in control trees was 250 micromol m(-3). No diurnal variation in [ABA(xyl)] was detected. With soil drying, [ABA(xyl)] increased to a maximum in summer (600 micromol m(-3)), but decreased again toward autumn; however, no significant increase in ABA flux to the leaves occurred. A decline in g(c) was detected when volumetric soil water content declined below 0.12. The decline in g(c) could not have been mediated by increasing [ABA(xyl)] because stomatal closure appeared to precede any increase in [ABA(xyl)]. Peak sap flow velocity data were used to estimate delivery times for root-to-shoot signals in 15-m tall trees. Under normal field conditions, a signal would take 12 days to travel from the site of production (roots) to the presumed site of action (shoots). However, under drought conditions it may take a chemical signal in excess of 6 weeks. We conclude that a feedforward model of short-term stomatal response to soil drying, based solely on the action of a chemical messenger from the roots, is not applicable in mature conifer trees because signal transmission is too slow.     

Genotypic variation in drought tolerance of poplar in relation to abscisic acid

We investigated effects of water stress and external abscisic acid (ABA) supply on shoot growth, stomatal conductance and water status in 1-year-old cuttings of a drought-sensitive poplar genotype Populus x euramericana cv. I-214 (Italica) and a drought-tolerant genotype P. 'popularis 35-44' (popularis). Populus popularis was more productive and maintained higher leaf water potentials throughout the drought treatment than cv. Italica. Supply of ABA to the xylem sap caused a greater decline in growth and more leaf abscission in shoots of cv. Italica than in shoots of P. popularis. Immediately after initiation of the drought treatment in P. popularis, the ABA concentration ([ABA]) of the xylem increased rapidly and stomatal conductance declined; however, stomatal conductance had returned to control values by the third day of the drought treatment, coincident with a gradual decline in xylem [ABA]. In contrast, xylem [ABA] of cv. Italica initially increased more slowly than that of P. popularis in response to the drought treatment, but the increase continued for 3 days at which time a tenfold increase in xylem [ABA] was observed that was followed by abscission of more than 40% of the leaves. We conclude that sensitivity of poplar roots to variation in soil water content varies by clone and that a rapid short-term accumulation of ABA in shoots in response to water stress may contribute to drought tolerance.     

Stomatal response of an anisohydric grapevine cultivar to evaporative demand, available soil moisture and abscisic acid

Stomatal responsiveness to evaporative demand (air vapour pressure deficit (VPD)) ranges widely between species and cultivars, and mechanisms for stomatal control in response to VPD remain obscure. The interaction of irrigation and soil moisture with VPD on stomatal conductance is particularly difficult to predict, but nevertheless is critical to instantaneous transpiration and vulnerability to desiccation. Stomatal sensitivity to VPD and soil moisture was investigated in Semillon, an anisohydric Vitis vinifera L. variety whose leaf water potential (Ψ(l)) is frequently lower than that of other grapevine varieties grown under similar conditions in the warm grape-growing regions of Australia. A survey of Semillon vines across seven vineyards revealed that, regardless of irrigation treatment, midday Ψ(l) was dependent on not only soil moisture but VPD at the time of measurement. Predawn Ψ(l) was more closely correlated to not only soil moisture in dry vineyards but to night-time VPD in drip-irrigated vineyards, with incomplete rehydration during high night-time VPD. Daytime stomatal conductance was low only under severe plant water deficits, induced by extremes in dry soil. Stomatal response to VPD was inconsistent across irrigation regime; however, in an unirrigated vineyard, stomatal sensitivity to VPD-the magnitude of stomatal response to VPD-was heightened under dry soils. It was also found that stomatal sensitivity was proportional to the magnitude of stomatal conductance at a reference VPD of 1kPa. Exogenous abscisic acid (ABA) applied to roots of Semillon vines growing in a hydroponic system induced stomatal closure and, in field vines, petiole xylem sap ABA concentrations rose throughout the morning and were higher in vines with low Ψ(l). These data indicate that despite high stomatal conductance of this anisohydric variety when grown in medium to high soil moisture, increased concentrations of ABA as a result of very limited soil moisture may augment stomatal responsiveness to low VPD.     

Abscisic acid accumulation in leaves of two contrasting hybrid poplar clones affected by nitrogen fertilization plus cyclic flooding and soil drying

Cuttings of hybrid Populus clones Tristis and Eugenei growing in pots in a greenhouse were treated with nitrogen fertilizer at two rates and subjected to repeated soil flooding or drying. Periodically, gas exchange measurements and radioimmunoassays, to determine abscisic acid (ABA) concentrations, were made on recently mature leaves.In both clones, photosynthesis and stomatal conductance were depressed five days after flooding, but leaf ABA concentrations remained relatively constant. In contrast, an initial, 9-day period of soil drying resulted in substantial ABA accumulation in leaves, which closely correlated with declines in photosynthesis and conductance. A second soil drying cycle of up to 9 days was less effective in modifying gas exchange and leaf ABA concentrations. High-N supply stimulated leaf ABA production as the soil dried. On the resumption of watering, gas exchange in Tristis recovered fully and rapidly and leaf ABA concentrations quickly returned to control values, whereas gas exchange in Eugenei recovered slowly and leaf ABA concentrations remained high for longer.Gas exchange in Eugenei was unaffected by soil drying until leaf ABA concentrations exceeded 100 ng g(dw) (-1), whereas Tristis showed a reduction in stomatal conductance and photosynthesis at leaf ABA concentrations of only 10 ng g(dw) (-1). A rise in internal CO(2) concentrations was associated with increased leaf ABA concentrations in Tristis, but not in Eugenei. Clonal differences in the relationship between gas exchange and leaf ABA concentration suggest contrasting physiological strategies for survival under prolonged drying conditions.     

Stomatal conductance of Acer rubrum ecotypes under varying soil and atmospheric water conditions: predicting stomatal responses with an abscisic acid-based model

A multiplicative model of stomatal conductance was developed and tested in two functionally distinct ecotypes of Acer rubrum L. (red maple). The model overcomes the main limitation of the commonly used Ball-Berry model (Ball et al. 1987) by accounting for stomatal behavior under soil drying conditions. We combined the Ball-Berry model with an integrated expression of abscisic acid (ABA)-based stomatal response to ABA concentration ([ABA]) in bulk leaf tissue (gfac), which coupled physiological changes at the leaf level with those in the root. The factor gfac = exp(-beta[ABA]L) incorporated the stomatal response to [ABA] into the Ball-Berry model by down regulating stomatal conductance (gs) in response to physiological changes in the root. The down regulation of gs is pertinent under conditions where soil drying may modify the delivery of chemical signals to leaf stomata. Model testing indicated that the multiplicative model was capable of predicting gs in red maple under wide ranges of soil and atmospheric conditions. Concordance correlation coefficients were high (between 0.59 and 0.94) for the tested ecotypes under three environmental conditions (atmospheric, rhizospheric and minimal stress). The study supported the use of gfac as a gas exchange function that controls water stress effects on gs and aids in the prediction of gs responses.     

Water-use efficiency in cork oak (Quercus suber) is modified by the interaction of water and light availabilities

We studied the interaction of light and water on water-use efficiency in cork oak (Quercus suber L.) seedlings. One-year-old cork oak seedlings were grown in pots in a factorial experiment with four light treatments (68, 50, 15 and 5% of full sunlight) and two irrigation regimes: well watered (WW) and moderate drought stress (WS). Leaf predawn water potential, which was measured at the end of each of two cycles, did not differ among the light treatments. Water-use efficiency, assessed by carbon isotope composition (delta(13)C), tended to increase with increasing irradiance. The trend was similar in the WW and WS treatments, though with lower delta(13)C in all light treatments in the WW irrigation regime. Specific leaf area increased with decreasing irradiance, and was inversely correlated with delta(13)C. Thus, changes in delta(13)C could be explained in part by light-induced modifications in leaf morphology. The relationship between stomatal conductance to water vapor and net photosynthesis on a leaf area basis confirmed that seedlings in higher irradiances maintained a higher rate of carbon uptake at a particular stomatal conductance, implying that shaded seedlings have a lower water-use efficiency that is unrelated to water availability.     

Hydraulic responses of whole vines and individual roots of kiwifruit (Actinidia chinensis) following root severance

Whole vine (K(plant)) and individual root (K(root)) hydraulic conductances were measured in kiwifruit (Actinidia chinensis Planch. var. chinensis 'Hort16A') vines to observe hydraulic responses following partial root system excision. Heat dissipation and compensation heat pulse techniques were used to measure sap flow in trunks and individual roots, respectively. Sap flux and measurements of xylem pressure potential (Ψ) were used to calculate K(plant) and K(root) in vines with zero and ～80% of roots severed. Whole vine transpiration (E), Ψ and K(plant) were significantly reduced within 24 h of root pruning, and did not recover within 6 weeks. Sap flux in intact roots increased within 24 h of root pruning, driven by an increase in the pressure gradient between the soil and canopy and without any change in root hydraulic conductance. Photosynthesis (A) and stomatal conductance (g(s)) were reduced, without significant effects on leaf internal CO(2) concentration (c(i)). Shoot growth rates were maintained; fruit growth and dry matter content were increased following pruning. The woody roots of kiwifruit did not demonstrate a rapid dynamic response to root system damage as has been observed previously in monocot seedlings. Increased sap flux in intact roots with no change in K(root) and only a moderate decline in shoot A suggests that under normal growing conditions root hydraulic conductance greatly exceeds requirements for adequate shoot hydration.     

Characterization of nitrogenous solutes in tissues and xylem sap of Leucaena leucocephala

Amino acid profiles of leaf, stem, and root tissues from nodulated and nonnodulated Leucaena leucocephala (Lam.) de Wit plants were determined by gas chromatography-mass spectrometry. High concentrations of mimosine and several other potentially toxic nonprotein amino acids, including pipecolic acid and two isomers of hydroxypipecolic acid, were identified in the tissues. Five metabolites remain unidentified. Of the foliar free amino acid nitrogen, 57-66% was associated with the potentially toxic amino acids. Major constituents in the leaf tissues of nonnodulated plants were mimosine and hydroxypipecolic acid (isomer 1). Mimosine was recovered in both the neutral plus basic and acidic amino acid fractions. Major differences between amino acid profiles of nodulated and nonnodulated roots were the low percentages of asparagine + aspartate (3.6% of the total pool compared to 33% in nodulated plants) and pipecolic acid in nonnodulated roots (1% of the total compared to 12.5% in nodulated plants). A novel plant betaine (dihydroxypipecolic acid betaine) was identified by fast-atom-bombardment mass spectrometry in leaf tissues, albeit at relatively low concentrations (< 1 micro mol per gram fresh weight). Analyses of the xylem sap collected from nodulated plants confirmed that Leucaena is an asparagine transporter, as suggested by the high concentrations of asparagine and the low concentrations of ureides in its root nodules. Amino acid profiles of xylem sap from nonnodulated plants showed extremely low concentrations of asparagine + aspartate (0.12 micro mol ml(-1)), whereas asparagine + aspartate was the major constituent (4.38 micro mol ml(-1)) in the xylem sap of nodulated plants. Two nonprotein amino acids, pipecolic acid and hydroxypipecolic acid, were major constituents of the xylem sap of nodulated and nonnodulated plants, respectively. Three unidentified compounds detected in xylem sap samples from both nodulated and nonnodulated plants did not correspond with any of the peaks characterized from tissue samples.     

Short-term effects of potassium fertilization on the hydraulic conductance of Laurus nobilis L

This study reports experimental evidence on the effect of short-term potassium fertilization on potassium uptake, tissue concentration and hydraulic conductance of pot-grown laurel plants. Potassium uptake and loading into the xylem of laurel seedlings increased within 24 h after fertilization. Potassium was not accumulated in roots and leaves, but the [K(+)] of xylem sap was 80% higher in fertilized plants (+K) than in potassium-starved plants (-K), as a likely result of recirculation between xylem and phloem. Increased xylem sap [K(+)] resulted in a 45% increase in transpiration rate, a 30% increase in plant hydraulic conductance (K(plant)) and a 120% increase in leaf-specific conductivity of the shoot (k(shoot)). We suggest that this increase was due to ion-mediated up-regulation of xylem hydraulics, possibly caused by the interaction of potassium ions with the pectic matrix of intervessel pits. The enhancement of hydraulic conductance following short-term potassium fertilization is a phenomenon that can be of advantage to plants for maintaining cell turgor, stomatal aperture and gas exchange rates under moderate drought stress. Our data provide additional support for the important role of potassium nutrition in agriculture and forestry.     

Water utilization,plant hydraulic properties and xylem vulnerability in three contrasting coffee (Coffea arabica) cultivars

Water use, hydraulic properties and xylem vulnerability to cavitation were studied in the coffee (Coffea arabica L.) cultivars San Ramon, Yellow Caturra and Typica growing in the field under similar environmental conditions. The cultivars differed in growth habit, crown morphology and total leaf surface area. Sap flow, stomatal conductance (g(s)), crown conductance (g(c)), apparent hydraulic conductance of the soil-leaf pathway (G(t)), leaf water potential (Psi(L)) and xylem vulnerability to loss of hydraulic conductivity were assessed under well-watered conditions and during a 21-day period when irrigation was withheld. Sap flow, g(c), and G(t) were greatest in Typica both with and without irrigation, lowest in San Ramon, which was relatively unresponsive to the withholding of irrigation, and intermediate in Yellow Caturra. The cultivars had similar g(s) when well watered, but withholding water decreased g(s) more in Typica and Yellow Caturra than in San Ramon. Typica had substantially lower Psi(L) near the end of the unirrigated period than the other cultivars (-2.5 versus -1.8 MPa), consistent with the relatively high sap flow in this cultivar. Xylem vulnerability curves indicated that Typica was less susceptible to loss of hydraulic conductivity than the other cultivars, consistent with the more negative Psi(L) values of Typica in the field during the period of low soil water availability. During soil drying, water use declined linearly with relative conductivity loss predicted from vulnerability curves. However, cultivar-specific relationships between water use and predicted conductivity loss were not observed because of pronounced hysteresis during recovery of water use following soil water recharge. All cultivars shared the same functional relationship between integrated daily sap flow and G(t), but they had different operating ranges. The three cultivars also shared common functional relationships between hydraulic architecture and water use despite consistent differences in water use under irrigated and dry soil conditions. We conclude that hydraulic architectural traits, rate of water use per plant and crown architecture are important determinants of short- and long-term variations in the water balance of Coffea arabica.     

Adaptations to soil drying in woody seedlings of African locust bean, (Parkia biglobosa (Jacq.) Benth.)

Stomatal conductance, transpiration and xylem pressure potential of African locust bean (Parkia biglobosa (Jacq.) Benth.) seedlings subjected from the sixth week after emergence to four weeks of continuous soil drought did not differ from those of well-watered, control plants until two-thirds of the available soil water had been used. In both well-watered and drought-treated plants, stomatal conductance was highest early in the day when vapor pressure deficits were low, but decreased sharply by midday when evaporative demand reached its highest value. There was no increase in stomatal conductance later in the day as vapor pressure deficit declined. The relationship between transpiration rate and xylem pressure potential showed non-linearity and hysteresis in both control and drought-treated plants, which seems to indicate that the plants had a substantial capacity to store water. The rate of leaf extension in African locust bean seedlings subjected to six consecutive 2-week cycles of soil drought declined relative to that of well-watered, control plants, whereas relative root extension increased. It appears that African locust bean seedlings minimized the impact of drought by: (1) restricting transpiration to the early part of the day when a high ratio of carbon gain to water loss can be achieved; (2) utilizing internally stored water during periods of rapid transpiration; (3) reducing the rate of leaf expansion and final leaf size in response to soil drought without reducing the rate of root extension, thereby reducing the ratio of transpiring leaf surface area to absorbing root surface area.     

Growth, nutrition and response to water stress of Pinus pinaster inoculated with ten dikaryotic strains of Pisolithus sp

Reconstituted dikaryons of Pisolithus sp. (Pers.) Coker & Couch from South Africa influenced growth parameters (shoot length, shoot/root ratio and leaf area), nutrition and physiological indicators (transpiration rate, stomatal conductance and xylem water potential) of maritime pine (Pinus pinaster Ait.) seedlings during drought and recovery from drought. Seedlings colonized with certain dikaryons were more sensitive to water stress and showed less mycorrhiza formation under water stress than seedlings colonized with other dikaryons. Control (uninoculated) seedlings were significantly smaller than those inoculated with dikaryons. Transpiration rate, stomatal conductance and xylem water potential varied among mycorrhizal treatments during the water stress and recovery periods. After rewatering, the controls and seedlings inoculated with dikaryon 34 x 20 had a weaker recovery in transpiration rate, stomatal conductance and xylem water potential than the other treatments and appeared to have experienced damage due to the water stress. Concentrations of various elements differed in the shoots of Pinus pinaster colonized by the various dikaryons. It is suggested that breeding of ectomycorrhizal fungi could constitute a new tool for improving reforestation success in arid and semi-arid zones.     

Effects of elevated carbon dioxide and drought on the growth and physiology of clonal Sitka spruce plants (Picea sitchensis (Bong.) Carr.)

Two-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.) plants from four clones were grown in naturally lit growth chambers for 6 months at either ambient (350 ppm) or ambient + 250 ppm (600 ppm) CO(2) concentration. Plants were grown in large boxes filled with peat, in a system that allowed the roots of individual plants to be harvested easily at the end of the growing season. Half of the boxes were kept well watered and half were allowed to dry out slowly over the summer. Plants growing in elevated CO(2) showed a 6.9% increase in mean relative growth rate compared to controls in the drought treatment and a 9.8% increase compared to controls in the well-watered treatment, though there was considerable variation in response among the different clones and water treatments. Rates of net CO(2) assimilation were higher and stomatal conductances were lower in plants grown in elevated CO(2) than in ambient CO(2) in both the well-watered and drought treatments. Both of these factors contributed to the doubling of instantaneous water use efficiency. The partitioning of biomass to roots was unaffected by elevated CO(2), but the ratio of needle mass/stems + branches mass decreased. Together with reduced stomatal conductance, this probably caused the observed increases in xylem pressure potentials with elevated CO(2).     

Abscisic acid in leaves and roots of willow: significance for stomatal conductance

Excised leaves and roots of willow (Salix dasyclados Wimm.) accumulated abscisic acid (ABA) in response to desiccation. The accumulation of ABA was greater in young leaves and roots than in old leaves and roots. In mature leaves, ABA accumulation was related to the severity and duration of the desiccation treatment. Water loss equal to 12% of initial fresh weight caused the ABA content of mature leaves to increase measurably within 30 min and to double in 2.5 h. The drying treatment caused significant (P = 0.05) reductions in leaf water potential and stomatal conductance. Recovery of leaf water potential to the control value occurred within 10 min of rewatering the dehydrated leaves, but recovery of stomatal conductance took an hour or longer, depending on the interval between dehydration and rewatering. The addition of ABA to the transpiration stream of well-watered excised leaves was sufficient to cause partial stomatal closure within 1 h and, depending on ABA concentration, more or less complete stomatal closure within 3 h. When the ABA solution was replaced with water, stomatal conductance increased at a rate inversely related to the concentration of the ABA solution with which the leaves had been supplied.     

Water stress responses of seedlings of four Mediterranean oak species

Effects of water stress on phenology, growth, stomatal activity and water status were assessed from April to November 1996 in 2-year-old seedlings of Quercus frainetto Ten. (Quercus conferta Kit.), Quercus pubescens Willd., Quercus macrolepis Kotschy (Quercus aegilops auct.) and Quercus ilex L. growing in containers in northern Greece. All four species developed more than 50% of their total leaf area before the beginning of June--an adaptation to arid climates. Well-irrigated plants tended to develop greater individual leaf area, number of leaves per plant, total plant leaf area, height and root:shoot ratios than water-stressed plants, but the difference between treatments was not significant for any parameter in any species. Quercus macrolepis appeared to be the most drought-tolerant of the four species. It maintained the highest number of leaves of the smallest size and increased the proportion of fine roots during drought. In all species, drought caused significant decreases in stomatal conductance and predawn and midday water potentials from mid-July until the end of August, when the lowest soil water content and highest mean daily air temperatures and midday leaf temperatures occurred; however, the responses were species-specific. Among the four species, Quercus macrolepis sustained the highest stomatal conductance despite very low water potentials, thus overcoming drought by means of desiccation tolerance. Quercus ilex decreased stomatal conductance even before severe water stress occurred, thereby avoiding desication during drought. Quercus pubescens had the highest water potential despite a high stomatal conductance, indicating that its leaf water status was independent of stomatal activity. Quercus frainetto was the least drought-resistant of the four species. During drought it developed very low water potentials despite markedly reduced stomatal aperture.