Influence of cadmium on water relations, stomatal resistance, and abscisic Acid content in expanding bean leaves

Ten day old bush bean plants (Phaseolus vulgaris L. cv Contender) were used to analyze the effects of 3 micromolar Cd on the time courses of expansion growth, dry weight, leaf water relations, stomatal resistance, and abscisic acid (ABA) levels in roots and leaves. Control and Cd-treated plants were grown for 144 hours in nutrient solution. Samples were taken at 24 hour intervals. At the 96 and 144 hour harvests, additional measurements were made on excised leaves which were allowed to dry for 2 hours. From the 48 hour harvest, Cd-treated plants showed lower leaf relative water contents and higher stomatal resistances than controls. At the same time, root and leaf expansion growth, but not dry weight, was significantly reduced. The turgor potentials of leaves from Cd-treated plants were nonsignificantly higher than those of control leaves. A significant increase (almost 400%) of the leaf ABA concentration was detected after 120 hours exposure to Cd. But Cd was found to inhibit ABA accumulation during drying of excised leaves. It is concluded that Cd-induced decrease of expansion growth is not due to turgor decrease. The possible mechanisms of Cd-induced stomatal closure are discussed.     

Stomatal response characteristics of Tradescantia virginiana grown at high relative air humidity

Plants produced at high relative air humidity (RH) show poor control of water loss after transferring to low RH, a phenomenon which is thought to be due to their stomatal behaviour. The stomatal anatomy and responses of moderate (55%) and high (90%) RH grown Tradescantia virginiana plants to treatments that normally induce stomatal closure, i.e. desiccation, abscisic acid (ABA) application and exposure to darkness were studied using attached or detached young, fully expanded leaves. Compared with plants grown at moderate RH the transpiration rate, stomatal conductance and aperture of high RH grown plants measured at the same condition (40% RH) were, respectively, 112, 139 and 132% in light and 141, 188 and 370% in darkness. Besides the differences in stomatal size (guard cell length was 56.7 and 73.3 µm for moderate and high RH grown plants, respectively), there was a clear difference in stomatal behaviour. The stomata responded to desiccation, ABA and darkness in both moderate and high RH grown plants, but the high variability of stomatal closure in high RH grown plants was striking. Some stomata developed at high RH closed in response to darkness or to a decrease in relative water content to the same extent as did stomata from moderate RH grown plants, whereas others closed only partly or did not close at all. Evidently, some as yet unidentified physiological or anatomical changes during development disrupt the normal functioning of some stomata in leaves grown at high RH. The failure of some stomata to close fully in response to ABA suggests that ABA deficiency was not responsible for the lack of stomatal closure in response to desiccation.     

Stomatal malfunctioning under low VPD conditions: induced by alterations in stomatal morphology and leaf anatomy or in the ABA signaling?

Exposing plants to low VPD reduces leaf capacity to maintain adequate water status thereafter. To find the impact of VPD on functioning of stomata, stomatal morphology and leaf anatomy, fava bean plants were grown at low (L, 0.23 kPa) or moderate (M, 1.17 kPa) VPDs and some plants that developed their leaves at moderate VPD were then transferred for 4 days to low VPD (M→L). Part of the M→L‐plants were sprayed with ABA (abscisic acid) during exposure to L. L‐plants showed bigger stomata, larger pore area, thinner leaves and less spongy cells compared with M‐plants. Stomatal morphology (except aperture) and leaf anatomy of the M→L‐plants were almost similar to the M‐plants, while their transpiration rate and stomatal conductance were identical to that of L‐plants. The stomatal response to ABA was lost in L‐plants, but also after 1‐day exposure of M‐plants to low VPD. The level of foliar ABA sharply decreased within 1‐day exposure to L, while the level of ABA‐GE (ABA‐glucose ester) was not affected. Spraying ABA during the exposure to L prevented loss of stomatal closing response thereafter. The effect of low VPD was largely depending on exposure time: the stomatal responsiveness to ABA was lost after 1‐day exposure to low VPD, while the responsiveness to desiccation was gradually lost during 4‐day exposure to low VPD. Leaf anatomical and stomatal morphological alterations due to low VPD were not the main cause of loss of stomatal closure response to closing stimuli.     

Model-based analysis of avoidance of ozone stress by stomatal closure in Siebold's beech (Fagus crenata)

Background and Aims

Resistance of plants to ozone stress can be classified as either avoidance or tolerance. Avoidance of ozone stress may be explained by decreased stomatal conductance during ozone exposure because stomata are the principal interface for entry of ozone into plants. In this study, a coupled photosynthesis–stomatal model was modified to test whether the presence of ozone can induce avoidance of ozone stress by stomatal closure.

Methods

The response of Siebold''s beech (Fagus crenata), a representative deciduous tree species, to ozone was studied in a free-air ozone exposure experiment in Japan. Photosynthesis and stomatal conductance were measured under ambient and elevated ozone. An optimization model of stomata involving water, CO₂ and ozone flux was tested using the leaf gas exchange data.

Key Results

The data suggest that there are two phases in the avoidance of ozone stress via stomatal closure for Siebold''s beech: (1) in early summer ozone influx is efficiently limited by a reduction in stomatal conductance, without any clear effect on photosynthetic capacity; and (2) in late summer and autumn the efficiency of ozone stress avoidance was decreased because the decrease in stomatal conductance was small and accompanied by an ozone-induced decline of photosynthetic capacity.

Conclusions

Ozone-induced stomatal closure in Siebold''s beech during early summer reduces ozone influx and allows the maximum photosynthetic capacity to be reached, but is not sufficient in older leaves to protect the photosynthetic system.     

Dynamics of adaptation of stomatal behaviour to moderate or high relative air humidity in Tradescantia virginiana

Chlorophyll fluorescence imaging was used to measure stomatalclosure in response to desiccation of Tradescantia virginianaleaves grown under high (90%) and moderate (55%) relative humidities(RHs), or transferred between these humidities. Stomata in leavesgrown at high RH were less responsive to desiccation than thoseof leaves grown at moderate RH. Stomata of plants transferredfrom moderate RH conditions to high RH showed the same diminishedclosure in response to desiccation as did stomata that developedat high RH. This response was found both when the leaves werefully expanded and when still actively expanding during themoderate RH pre-treatment. Four days of exposure to high RHwas the minimal exposure time to induce the diminished closureresponse. When leaves were grown in high RH prior to a 10 dmoderate RH treatment, the reduced stomatal closure responseto desiccation was only reversed in leaves (regions) which wereactively expanding during moderate RH treatment. This indicatesthat with respect to stomatal responses to desiccation, highRH leaf regions have a limited capacity to adapt to moderateRH conditions. The decrease in responsiveness to desiccationof the stomata, induced by long-term exposure to high RH, wasnot due to osmotic adjustment in the leaves. Within 1 d aftertransferring moderate RH-grown plants to a high RH, the abscisicacid (ABA) concentration of their leaves decreased to the lowlevel of ABA found in high RH-grown leaves. The closure responsein leaves exposed to high RH for 5 d, however, could not befully restored by the application of ABA. Transferring plantsfrom high to moderate RH resulted in increased ABA levels within2 d without a recovery of the stomatal closing response. Itis discussed that the diminished stomatal closure in plantsexposed to high RH could be due to changes in the signallingpathway for ABA-related closure of stomata or to an increasedsequestration of ABA by mesophyll tissue or the symplast inthe epidermis, induced by a longer period (several days) ofa low ABA level. Key words: Abscisic acid, desiccation, PSII efficiency, relative water content, stomatal closure, vapour pressure deficit, water potential Received 8 October 2007; Revised 5 November 2007 Accepted 9 November 2007     

Water Stress Reduces Ozone Injury via a Stomatal Mechanism

Various studies have shown that water-stressed plants are more tolerant of ozone exposures than are unstressed plants. Two probable explanations for this tolerance are (a) stomatal closure which reduces ozone uptake and (b) biochemical or anatomical changes within the leaves. Phaseolus vulgaris cv Pinto bean plants were established and transferred to membrane systems which controlled the osmotic potential around the roots at −35 or −80 kilopascals for 5 days prior to ozone treatment (0 or 1.0 microliters per liter for 2 hours). Both water-stressed and unstressed plants were sprayed with various concentrations of abscisic acid to close the stomata or with fusicoccin to induce stomata opening. The abaxial stomatal resistances of primary and trifoliate leaves were measured just prior to ozone exposure. Plant response to ozone was determined by stress ethylene production and chlorophyll loss. Both water stress and abscisic acid induced stomatal closure and reduced ozone injury. In water-stressed plants, fusicoccin induced stomatal opening and those plants were as sensitive to ozone as were the non-water-stressed plants. These data suggest that water stress protects plants from ozone injury mainly through its influence on stomatal aperture rather than through biochemical or anatomical changes.     

Avoiding high relative air humidity during critical stages of leaf ontogeny is decisive for stomatal functioning

Plants of several species, if grown at high relative air humidity (RH ≥85%), develop stomata that fail to close fully in case of low leaf water potential. We studied the effect of a reciprocal change in RH, at different stages of leaf expansion of Rosa hybrida grown at moderate (60%) or high (95%) RH, on the stomatal closing ability. This was assessed by measuring the leaf transpiration rate in response to desiccation once the leaves had fully expanded. For leaves that started expanding at high RH but completed their expansion after transfer to moderate RH, the earlier this switch took place the better the stomatal functioning. Leaves initially expanding at moderate RH and transferred to high RH exhibited poor stomatal functioning, even when this transfer occurred very late during leaf expansion. Applying a daily abscisic acid (ABA) solution to the leaves of plants grown at continuous high RH was effective in inducing stomatal closure at low water potential, if done before full leaf expansion (FLE). After FLE, stomatal functioning was no longer affected either by the RH or ABA level. The results indicate that the degree of stomatal adaptation depends on both the timing and duration of exposure to high RH. It is concluded that stomatal functionality is strongly dependent on the humidity at which the leaf completed its expansion. The data also show that the effect of ambient RH and the alleviating role of ABA are restricted to the period of leaf expansion.     

Dynamics of spatial heterogeneity of stomatal closure in Tradescantia virginiana altered by growth at high relative air humidity

The spatial heterogeneity of stomatal closure in response to rapid desiccation of excised well-watered Tradescantia virginiana leaves grown at moderate (55%) or high (90%) relative air humidity (RH) was studied using a chlorophyll fluorescence imaging system under non-photorespiratory conditions. Following rapid desiccation, excised leaves grown at high RH had both a greater heterogeneity and a higher average value of PSII efficiency (Phi(PSII)) compared with leaves grown at moderate RH. Larger decreases in relative water content resulted in smaller decreases in water potential and Phi(PSII) of high RH-grown leaves compared with moderate RH-grown leaves. Moreover, the Phi(PSII) of excised high RH-grown leaves decreased less with decreasing water potential, implying that the stomata of high RH-grown leaves are less sensitive to decreases in leaf water potential compared with moderate RH-grown leaves. After desiccation, some non-closing stomata were distributed around the main vein in high RH-grown leaves. Direct measurements of stomatal aperture showed 77% stomatal closure in the margins after 2 h desiccation compared with 40% closure of stomata in the main-vein areas in high RH-grown leaves. Faster closure of stomata in leaf margins compared with main-vein areas of leaves grown at high RH was related to substantially lower relative water content in these areas of the leaves.     

Foliar abscisic acid content underlies genotypic variation in stomatal responsiveness after growth at high relative air humidity

Background and Aims

Stomata formed at high relative air humidity (RH) respond less to abscisic acid (ABA), an effect that varies widely between cultivars. This study tested the hypotheses that this genotypic variation in stomatal responsiveness originates from differential impairment in intermediates of the ABA signalling pathway during closure and differences in leaf ABA concentration during growth.

Methods

Stomatal anatomical features and stomatal responsiveness to desiccation, feeding with ABA, three transduction elements of its signalling pathway (H₂O₂, NO, Ca²⁺) and elicitors of these elements were determined in four rose cultivars grown at moderate (60 %) and high (90 %) RH. Leaf ABA concentration was assessed throughout the photoperiod and following mild desiccation (10 % leaf weight loss).

Key Results

Stomatal responsiveness to desiccation and ABA feeding was little affected by high RH in two cultivars, whereas it was considerably attenuated in two other cultivars (thus termed sensitive). Leaf ABA concentration was lower in plants grown at high RH, an effect that was more pronounced in the sensitive cultivars. Mild desiccation triggered an increase in leaf ABA concentration and equalized differences between leaves grown at moderate and high RH. High RH impaired stomatal responses to all transduction elements, but cultivar differences were not observed.

Conclusions

High RH resulted in decreased leaf ABA concentration during growth as a result of lack of water deficit, since desiccation induced ABA accumulation. Sensitive cultivars underwent a larger decrease in leaf ABA concentration rather than having a higher ABA concentration threshold for inducing stomatal functioning. However, cultivar differences in stomatal closure following ABA feeding were not apparent in response to H₂O₂ and downstream elements, indicating that signalling events prior to H₂O₂ generation are involved in the observed genotypic variation.     

A Negative Hydraulic Message from Oxygen-Deficient Roots of Tomato Plants? (Influence of Soil Flooding on Leaf Water Potential,Leaf Expansion,and Synchrony between Stomatal Conductance and Root Hydraulic Conductivity)

Four to 10 h of soil flooding delayed and suppressed the normal daily increase in root hydraulic conductance (Lp) in tomato (Lycopersicon esculentum Mill. cv Ailsa Craig) plants. The resulting short-term loss of synchrony between Lp and stomatal conductance decreased leaf water potential ([psi]L) relative to well-drained plants within 2 h. A decrease in [psi]L persisted for 8 h and was mirrored by decreased leaf thickness measured using linear displacement transducers. After 10 h of flooding, further closing of stomata and re-convergence of Lp in flooded and well-drained roots returned [psi]L to control values. In the second photoperiod, Lp in flooded plants exceeded that in well-drained plants in association with much increased Lp and decreased stomatal conductance. Pneumatic balancing pressure applied to roots of intact flooded plants to prevent temporary loss of [psi]L in the 1st d did not modify the patterns of stomatal closure or leaf expansion. Thus, the magnitude of the early negative hydraulic message was neither sufficient nor necessary to promote stomatal closure and inhibit leaf growth in flooded tomato plants. Chemical messages are presumed to be responsible for these early responses to soil flooding.     

Physiology of Polyploid Plants: Water Balance in Autotetraploid and Diploid Tomato under Low and High Salinity

Diploid and autotetraploid plants of the cultivated tomato Lycopersicon esculentum cv. Lukullus (Luk) were studied under low and high salinity. Polyploids had a higher water content than diploid plants. Water content in both plant types decreased under salinity, the decrease being smaller in the polyploid plants. Dry weight of whole young plants decreased in both diploid and polyploid plants under salinity, the decrease being smaller in the latter. Transpiration of whole plants, grown in control solution, was lower in polyploid than in diploid plants and decreased more under salinity in the latter. Rate of change of water loss of detached drying leaves was similar in diploid and polyploid plants. Leaves of control diploid plants, however, lost more water per unit leaf area during the phase of stomatal closure apparently due to higher stomatal density. Polyploid plants had fewer but more open stomata per unit leaf area, under both control and saline conditions. Root pressure, determined only under control conditions, seemed to be higher in polyploid plants. No difference in Cl^? concentration per unit leaf dry weight was found between diploid and polyploid plants grown in either control or NaCl solution.     

Some physiological and growth responses of kiwi fruit (Actinidia chinensis) to flooding

The effects of long-term flooding on the growth of six-month-old Actinidia chinensis Planch cv. Abbot plants and some effects on stomatal behaviour and leaf water relations were examined under controlled conditions for 28 days. Flooding caused stomatal closure and decreases in transpiration rate, xylem water potential, osmotic potential and turgor potential. Flooding also caused inhibition of the dry weight increase of leaves plus stems and of roots, chlorosis and necrosis of leaves, production of hypertrophied lenticels and the appearance of a small number of adventitious roots on the submerged portions of the stems. Rapid and partial stomatal closure by flooding may not only be due to the passive mechanical response which follows leaf dehydration, since flooded plants showed an increase in xylem water potential and osmotic potential during the first days of the experiment. The marked intolerance of Actinidia chinensis to flooding has been a serious barrier to its culture in poorly drained soils, hence careful irrigation management is required.     

Stomatal closure during leaf dehydration,correlation with other leaf physiological traits

The question as to what triggers stomatal closure during leaf desiccation remains controversial. This paper examines characteristics of the vascular and photosynthetic functions of the leaf to determine which responds most similarly to stomata during desiccation. Leaf hydraulic conductance (K(leaf)) was measured from the relaxation kinetics of leaf water potential (Psi(l)), and a novel application of this technique allowed the response of K(leaf) to Psi(l) to be determined. These "vulnerability curves" show that K(leaf) is highly sensitive to Psi(l) and that the response of stomatal conductance to Psi(l) is closely correlated with the response of K(leaf) to Psi(l). The turgor loss point of leaves was also correlated with K(leaf) and stomatal closure, whereas the decline in PSII quantum yield during leaf drying occurred at a lower Psi(l) than stomatal closure. These results indicate that stomatal closure is primarily coordinated with K(leaf). However, the close proximity of Psi(l) at initial stomatal closure and initial loss of K(leaf) suggest that partial loss of K(leaf) might occur regularly, presumably necessitating repair of embolisms.     

The effects of ozone on growth and stomatal response in the F<Subscript>2</Subscript> generation of hybrid poplar (<Emphasis Type="Italic">Populus trichocarpa × Populus deltoides</Emphasis>)

Thirty-six F₂ hybrid poplar (Populus trichocarpa × P. deltoides) clones were fumigated with ozone to record its effects on growth, correlate them with stomatal response and screen for ozone sensitivity. Fumigation was applied for 6 to 9 h each day for approximately 3 months at ozone concentrations of 85 to 128 μg g⁻¹ using open-top chambers. Height, diameter, number of leaves, stomatal conductance, transpiration rate, total biomass, biomass components and root/shoot ratios were reduced by ozone stress. Percent of leaf fall in ozone-treated plants was nearly three times higher than in control plants exposed to charcoal-filtered air. Leaf senescence, because of ozone exposure, did not appear to be associated with reduced biomass production. Some clones had a high percentage of leaf-fall with ozone exposure, but were able to maintain total biomass production near that of the control. Their response may be an example of an ability to adjust or compensate for ozone damage. There was no significant or consistent relationship between stomatal conductance and total biomass or the change in stomatal conductance as a result of ozone exposure and the change in total biomass. Taken together, these results suggest that effects of ozone on poplar growth cannot be solely correlated to changes in stomatal conductance, more physiological and biochemical parameters should be examined.     

The role of abscisic acid in disturbed stomatal response characteristics of Tradescantia virginiana during growth at high relative air humidity

In this study, the role of abscisic acid (ABA) in altered stomatal responses of Tradescantia virginiana leaves grown at high relative air humidity (RH) was investigated. A lower ABA concentration was found in leaves grown at high RH compared with leaves grown at moderate RH. As a result of a daily application of 20 microM ABA to leaves for 3 weeks during growth at high RH, the stomata of ABA-treated leaves grown at high RH showed the same behaviour as did the stomata of leaves grown at moderate RH. For example, they closed rapidly when exposed to desiccation. Providing a high RH around a single leaf of a plant during growth at moderate RH changed the stomatal responses of this leaf. The stomata in this leaf grown at high RH did not close completely in response to desiccation in contrast to the stomata of the other leaves from the same plant. The ABA concentration on a fresh weight basis, though not on a dry weight basis, of this leaf was significantly lower than that of the others. Moreover, less closure of stomata was found in the older leaves of plants grown at high RH in response to desiccation compared with younger leaves. This was correlated with a lower ABA concentration in these leaves on a fresh weight basis, though not on a dry weight basis. Stomata of leaves grown at moderate RH closed in response to short-term application of ABA or sodium nitroprusside (SNP), while for leaves grown at high RH there was a clear difference in stomatal responses between the leaf margins and main-vein areas. The stomatal aperture in response to short-term application of ABA or SNP at the leaf margins of leaves grown at high RH remained significantly wider than in the main-vein areas. It was concluded that: (i) a long-term low ABA concentration in well-watered plants during growth at high RH could be a reason for less or no stomatal closure under conditions of drought stress; and (ii) the long-term ABA concentration on a fresh weight basis rather than on a dry weight basis is likely to be responsible for structural or physiological changes in stomata during leaf growth.     

A comprehensive analysis of the physiological and anatomical components involved in higher water loss rates after leaf development at high humidity

To better understand the poor regulation of water loss after leaf development at high relative air humidity (RH), the relative importance of the physiological and anatomical components was analyzed focusing on cultivars with a contrasting sensitivity to elevated RH. The stomatal responsiveness to three closing stimuli (desiccation, abscisic acid feeding, light/dark transition), as well as several stomatal features (density, index, size and pore dimensions) and the cuticular transpiration rate (CTR) were determined in four rose cultivars, grown under moderate (60%) and high (95%) RH. Moreover, the effects of changes in stomatal density and pore dimensions on the stomatal conductance (g_s) were quantified using a modified version of the Brown and Escombe equation. Higher water loss, as a result of plant growth at high RH, was primarily caused by an increase in residual g_s, and to a lesser extent due to higher CTR. It was estimated that in leaflets subjected to desiccation the enhanced g_s in high RH- as compared to moderate RH-grown plants was mostly due to poor stomatal functionality and to a lesser extent the combined result of higher stomatal density and longer pore length. It is concluded that the reduced degree and, specially, the reduced rate of stomatal closure are the primary causes of the large genotypic variation in the control of water loss in high RH-grown plants. Furthermore, it was found that although changes in stomatal length have no influence on stomatal functionality, changed anatomical features per se represent a significant and direct contribution to the increased water loss.     

Effects of leaf wetting and high humidity on stomatal function in leafy cuttings and intact plants of Corylus maxima

When rooted cuttings of Corylus maxima Mill. cv. Purpurea are moved from the wet and humid conditions of the rooting environment, the leaves frequently shrivel and die. Since the newly formed adventitious root system has been shown to be functional in supplying water to the shoot, stomatal behaviour in C. maxima was investigated in relation to the failure to prevent desiccation. Stomatal conductance (g_s) in expanding leaves (L3) of cuttings increased almost 10-fold over the first 14 days in the rooting environment (fog), from 70 to 650 mmol m⁻² s⁻¹. In contrast, g_s of expanded leaves (L1) changed little and was in the region of 300 mmol m⁻² s⁻¹. Midday leaf water potential was much higher in cuttings than in leaves on the mother stock-plant (−0.5 versus −1.2 MPa) even before any roots were visible. Despite this, leaf expansion of L3 was inhibited by >50% in cuttings and stomata showed a gradual reduction in their ability to close in response to abscisic acid (ABA). To determine whether the loss of stomatal function in cuttings was due to severance or to unnaturally low vapour pressure deficit and wetting in fog, intact plants were placed alongside cuttings in the rooting environment. The intact plants displayed reductions in leaf expansion and in the ability of stomata to close in response to dark, desiccation and ABA. However, in cuttings, the additional effect of severance resulted in smaller leaves than in intact plants and more severe reduction in stomatal closure, which was associated with a 2.5-fold increase in stomatal density and distinctively rounded stomatal pores. The similarities between stomatal dysfunction in C. maxima and that observed in many species propagated in vitro are discussed, as is the possible mechanism of dysfunction.     

Interaction of water supply and N in wheat

The purpose of this study was to investigate effects of N nutrition and water stress on stomatal behavior and CO₂ exchange rate in wheat (Triticum aestivum L. cv Olaf). Wheat plants were grown hydroponically with high (100 milligrams per liter) and low (10 milligrams per liter) N. When plants were 38 days old, a 24-day water stress cycle was begun. A gradual increase in nutrient solution osmotic pressure from 0.03 to 1.95 mega Pascals was achieved by incremental additions of PEG-6,000. Plants in both N treatments adjusted osmotically, although leaf water potential was consistently lower and relative water content greater for low N plants in the first half of the stress cycle. Leaf conductance of high N plants appeared greater than that of low N plants at high water potentials, but showed greater sensitivity to reductions in water potential as indicated by earlier stomatal closure during the stress cycle. The apparent greater stomatal sensitivity of high N plants was associated with a curvilinear relationship between leaf conductance and leaf water potential; low N plants exhibited more of a threshold response. Trends in [CO₂]_INT throughout the stress cycle indicated nonstomatal effects of water stress on CO₂ exchange rate were greater in high N plants. Although estimates of [CO₂]_INT were generally lower in high N plants, they were relatively insensitive to leaf water potential-induced changes in leaf conductance. In contrast, [CO₂]_INT of low N plants dropped concomitantly with leaf conductance at low leaf water potentials. Oxygen response of CO₂ exchange rate for both treatments was affected less by reductions in water potential than was CO₂ exchange rate at 2.5% O₂, suggesting that CO₂ assimilation capacity of the leaves was affected more by reductions in leaf water potential than were processes related to photorespiration.     

Early stomatal closure in waterlogged pea plants is mediated by abscisic acid in the absence of foliar water deficits

Abstract Soil waterlogging decreased leaf conductance (interpreted as stomatal closure) of vegetative pea plants (Pisuin sativum L. cv. ‘Sprite’) approximately 24 h after the start of flooding, i.e. from the beginning of the second 16 h-long photo-period. Both adaxial and abaxial surfaces of leaves of various ages and the stipules were affected. Stomatal closure was sustained for at least 3 d with no decrease in foliar hydration measured as water content per unit area, leaf water potential or leaf water saturation deficit. Instead, leaves became increasingly hydrated in association with slower transpiration. These changes in the waterlogged plants over 3 d were accompanied by up to 10-fold increases in the concentration of endogenous abscisic acid (ABA). Waterlogging also increased foliar hydration and ABA concentrations in the dark. Leaves detached from non-waterlogged plants and maintained in vials of water for up to 3 d behaved in a similar way to leaves on flooded plants, i.e. stomata closed in the absence of a water deficit but in association with increased ABA content. Applying ABA through the transpiration stream to freshly detached leaflets partially closed stomata within 15 min. The extractable concentrations of ABA associated with this closure were similar to those found in flooded plants. When an ABA-deficient ‘wilty’ mutant of pea was waterlogged, the extent of stomatal closure was less pronounced than that in ordinary non-mutant plants, and the associated increase in foliar ABA was correspondingly smaller. Similarly, waterlogging closed stomata of tomato plants within 24 h, but no such closure was seen in ‘flacca’, a corresponding ABA-deficient mutant. The results provide an example of stomatal closure brought about by stress in the root environment in the absence of water deficiency. The correlative factor operating between the roots and shoots appeared to be an inhibition of ABA transport out of the shoots of flooded plants, causing the hormone to accumulate in the leaves.     

Leaf Age as a Determinant in Stomatal Control of Water Loss from Cotton during Water Stress

The stomatal resistance of individual leaves of young cotton plants (Gossypium hirsutum L. var. Stoneville 213) was measured during a period of soil moisture stress under conditions of constant evaporative demand. When plants were subjected to increasing soil water stress, increases in stomatal resistance occurred first on the lower leaves and the stomata on the upper surfaces were the most sensitive to decreasing leaf-water potential. Stomatal closure proceeded from the oldest leaves to the youngest as the stress became more severe. This apparent effect of leaf age was not due to radiation differences during the stress period. Radiation adjustments on individual leaves during their development altered the stomatal closure potential for all leaves, but did not change the within-plant pattern. Our data indicate that no single value of leaf water potential will adequately represent a threshold for stomatal closure in cotton. Rather, the stomatal resistance of each leaf is uniquely related to its own water potential as modified by age and radiation regime during development. The effect of age on stress-induced stomatal closure was not associated with a loss of potassium from older leaves. Increases in both the free and bound forms of abscisic acid were observed in water-stressed plants, but the largest accumulations occurred in the youngest leaves. Thus, the pattern of abscisic acid accumulation in response to water stress did not parallel the pattern of stomatal closure induced by water stress.