Changes in leaf gas exchange, water relations, biomass production and solute accumulation in Phragmites australis under hypoxic conditions

The physiological responses to hypoxic stress were studied in the common reed, Phragmites australis (Cav.) Trin. ex Steudel. Growth, leaf gas exchange, water (and ion) relations and osmotic adjustment were determined in hydroponically grown plants exposed to 10, 20 and 30 days of oxygen deficiency. The highest growth of reed seedlings was found in normoxic (aerobic) conditions. Treatment effects on biomass production were relatively consistent within each harvest. Leaf water potential and osmotic potential declined significantly as hypoxia periods increased. However, leaf turgor pressure showed a consistent pattern of increase, suggesting that reed plants adjusted their water status by osmotic adjustment in response to root hypoxia. After 20 and 30 days in the low oxygen treatment, net CO₂ assimilation and stomatal conductance were positively associated and the former variable also had a strong positive relationship with transpiration. Short-term hypoxic stress had a slight effect on the ionic status (K⁺, Ca²⁺ and Mg²⁺) of reed plants. In contrast, soluble sugar concentrations increased more under hypoxic conditions as compared to normoxia. These findings indicate that hypoxia slightly affected the physiological behavior of reed plants.     

Combined effect of NaCl-salinity and hypoxia on growth,photosynthesis, water relations and solute accumulation in Phragmites australis plants

Aim of the present study was to investigate the effects of two key environmental factors of estuarine ecosystems, salinity and hypoxia, on the physiological attributes in reed plants (Phragmites australis (Cav.) Trin. ex Steudel). Growth, leaf gas exchange, water (and ion) relations, and osmotic adjustment were determined in hydroponically grown plants exposed to hypoxia at varying NaCl-salinity concentrations (0, 50, 100, and 200 mM). Plants grew well under hypoxia treatment with standard nutrient solution without added salt and at NaCl concentrations up to 100 mM. Reed plants were able to produce and allocate phytomass to all their organs even at the highest salt level (200 mM NaCl). In plants subjected to hypoxia at various water potentials no clear relationships were found between growth and photosynthetic parameters except for g_s, whereas growth displayed a highly significant correlation with plant–water relations. A and g_s of reed plants treated with hypoxia at varying water potential of nutrient solutions were positively correlated and the former variable also had a strong positive relationship with E. Leaf Ψ_w and Ψ_π followed a similar trend and declined significantly as water potential of watering solutions was lowered. Highly significant positive correlations were identified between leaf Ψ_w and photosynthetic parameters. At all NaCl concentrations, the increase in total inorganic ions resulted from increased Na⁺ and Cl⁻ while K⁺, Ca²⁺, and Mg²⁺ concentrations decreased with increasing osmolality of nutrient solutions. Common reed has an efficient mechanism of Na⁺ exclusion from the leaves and exhibited a high leaf K⁺/Na⁺ selectivity ratio over a wide range of salinities under hypoxia treatment. In Phragmites australis grown in 200 mM NaCl, K⁺ contributed 17% toΨ_π, whereas Na⁺ and Cl⁻ accounted for only 11% and 6%, respectively. At the same NaCl concentration, the estimated contribution of proline to Ψ_π was less than 0.2%. Changes in leaf turgor occurred with a combined effect of salinity and hypoxia, suggesting that reed plants could adjust their water status sufficiently.     

Influence of NaCl-salinity on growth,photosynthesis, water relations and solute accumulation in <Emphasis Type="Italic">Phragmites australis</Emphasis>

The aim of this study was to investigate the effects of NaCl-salinity on the physiological attributes in common reed, Phragmites australis (Cav.) Trin. ex Steudel. Plants grew optimally under salinity treatment with standard nutrient solution without added salt and at NaCl concentrations up to 100 mM. Applied for 21 days, NaCl-salinity (300 and 500 mM) caused a significant reduction in growth allocation of all different tissues of P. australis. Shoot growth of reed plants displayed a highly significant correlation with plant–water relations and photosynthetic parameters. The net photosynthetic rate and stomatal conductance of reed plants treated with NaCl-salinity at varying osmotic potential (ψ_π) of nutrient solutions were positively correlated, and the former variable also had a strong positive relationship with transpiration rate. Leaf water potential and ψ_π followed similar trends and declined significantly as ψ_π of watering solutions was lowered. The increase in total inorganic nutrients resulting from increased Na⁺ and Cl⁻ in all tissues and K⁺, Ca²⁺ and Mg²⁺ concentrations were maintained even at the most extreme salt concentration. Common reed exhibited high K⁺/Na⁺ and Ca²⁺/Na⁺ selectivity ratios over a wide range of salinities under NaCl-salinity. These findings suggest that reed plants were able to adapt well to high salinities by lowering their leaf ψ_π and the adjustment of osmotically active solutes in the leaves.     

Responses to drought preconditioning in<Emphasis Type="Italic"> Eucalyptus globulus</Emphasis> Labill. provenances

Shoot water relations and morphological responses to drought preconditioning were studied by subjecting 5-month-old seedlings of three provenances of Eucalyptus globulus to different water regimes for 36 days in a greenhouse pot study. Moderately stressed plants were watered every 6 days and severely stressed plants were watered every 9 days. Control plants were watered daily. Drought cycles induced significant changes in morphological and physiological characteristics. Preconditioned seedlings were smaller in size, root collar diameter, height, and leaf area than control seedlings. Shoot/root ratio was not affected by drought. Osmotic potential at full turgor (ψπ_FT) and osmotic potential at turgor loss point (ψπ_TLP) were significantly lower and the magnitude of osmotic adjustment was significantly higher under the severe than under the moderate stress treatment. In severely stressed plants a decrease of turgid mass/dry mass contributed to osmotic adjustment. In a subsequent acclimation test, preconditioned seedlings showed higher values of stomatal conductance, predawn relative water content and water potential and lower mortality than control plants. These variables were significantly related to ψπ_FT. We assume that the reduced leaf area and osmotic adjustment observed in preconditioned seedlings contributed to drought acclimation in the selected E. globulus provenances leading to better rates of gas exchange and improved water status than non-conditioned plants. Provenances exhibited differences in their responses to drought, albeit mainly morphological differences. E. globulus subsp. bicostata from Tumbarumba grew more quickly (larger diameter and height relative growth rate) than the other provenances, implying a greater ability to tolerate water stress. It can be expected that preconditioned seedlings will display greater tolerance of water stress than non-conditioned plants and perform better during early establishment (higher survival and early growth).     

Effects of 24‐epibrassinolide on plant growth,osmotic regulation and ion homeostasis of salt‐stressed canola

This study evaluated effects of foliar spraying 24‐epibrassinoide (24‐EBL) on the growth of salt‐stressed canola. Seedlings at the four‐leaf stage were treated with 150 mm NaCl and different concentrations of 24‐EBL (10^?6, 10^?8, 10^?10, 10^?12 m ) for 15 days. A concentration of 10^?10 m 24‐EBL was chosen as optimal and used in a subsequent experiment on plant biomass and leaf water potential parameters. The results showed that 24‐EBL mainly promoted shoot growth of salt‐stressed plants and also ameliorated leaf water status. Foliar spraying of salt‐stressed canola with 24‐EBL increased osmotic adjustment ability in all organs, especially in younger leaves and roots. This was mainly due to an increase of free amino acid content in upper leaves, soluble sugars in middle leaves, organic acids and proline in lower leaves, all of these compounds in roots, as well as essential inorganic ions. Na⁺ and Cl^? sharply increased in different organs under salt stress, and 24‐EBL reduced their accumulation. 24‐EBL improved the uptake of K⁺, Ca²⁺, Mg²⁺ and NO₃^? in roots, which were mainly transported to upper leaves, while NO₃^? was mainly transported to middle leaves. Thus, 24‐EBL improvements in ion homeostasis of K⁺/Na⁺, Ca²⁺/Na⁺, Mg²⁺/Na⁺ and NO₃^?/Cl^?, especially in younger leaves and roots, could be explained. As most important parts, younger leaves and roots were the main organs protected by 24‐EBL via improvement in osmotic adjustment ability and ion homeostasis. Further, physiological status of growth of salt‐stressed canola was ameliorated after 24‐EBL treatment.     

Does Cytokinin Transport from Root-To-Shoot in the Xylem Sap Regulate Leaf Responses to Root Hypoxia?

We have examined the hypothesis that cytokinins transportedfrom roots to shoots affects leaf growth, stomatal conductance,and cytokinin concentration of leaves of Phaseolus and a hybridpoplar (Populus trichocarpa x Populus deltoides) with hypoxicroots. Because cytokinins may interact with other substances,potassium and calcium concentrations were determined in xylemsap of Populus plants with hypoxic and aerated roots while gibberellin(GA) concentrations were measured in shoot tissues. Root hypoxiadecreased leaf growth and closed stomata in both species. Inboth species, fluxes of cytokinins out of the roots were reduced,but no differences in bulk leaf concentrations were measuredbetween the hypoxic and aerated plants. Shoots with aeratedroots contained slightly higher concentrations of GA₁ and GA₃than shoots from hypoxic plants. There were no differences incalcium or potassium concentrations in xylem sap between aerationtreatments. Exogenously applied cytokinins did not alleviatethe growth or stomatal responses caused by root hypoxia. Informationon the site(s) and mechanism(s) of cytokinin action and theways in which cytokinins are compartmentalized within plantcells will be required to understand the physiological significanceof cytokinin transport in the transpirational stream. Key words: Cytokinins, hypoxia, Populus, Phaseolus     

Role of Osmotic Potential Gradients during Water Stress and Leaf Senescence in Fragaria virginiana

The physiological basis underlying differences in sensitivity of different aged leaves to water stress was investigated in Fragaria virginiana Duchesne. Differential susceptibility of only older leaves to water stress in the field during summer months appeared related to gradients in leaf osmotic potential within the plant and by an age dependency in the ability of leaves to adjust osmotically when challenged by periodic water deficits. Under greenhouse conditions, older leaves senesced invariably during an imposed water stress while control leaves of comparable age and stressed younger leaves remained green. Osmotic potentials of intermediate aged and younger leaves became approximately 1 to 2 bars lower after a single cycle of imposed stress and up to 10 bars lower after two cycles of stress. Pronounced gradients in leaf osmotic potential within individual whole plants were observed following two cycles of water stress that were significantly different from control values. Osmotic adjustment was dependent on leaf age with the greatest capacity for adjustment in the intermediate aged leaves. Loss of osmotic adjustment was rapid upon rewatering with a half-life of 4 days. An irreversible component of adjustment was observed, amounting to about 10% (or 2 bars) of the maximally adjusted state. This irreversible component could be accounted for in part by significant changes in cell size and other anatomical alterations in the leaf that affect cellular osmotic volume, and, hence, cellular water relations.     

Effects of nitrogen nutrition and root medium water potential on growth, nitrogen uptake and osmotic adjustment of rice

The effects of nitrogen (N) nutrition on growth, N uptake and leaf osmotic potential of rice plants (Oryza sativa L. ev. IR 36) during simulated water stress were determined. Twenty-one-day-old seedlings in high (28.6 × 10 ^?4M) and low (7.14 × 10 ⁴M) N levels were exposed to decreased nutrient solution water potentials by addition of polyethylene glycol 6000. The roots were separated from the solution by a semi-permeable membrane. Nutrient solution water potential was ?0.6 × 10⁵ Pa and was lowered stepwise to ?1 × 10⁵, ?2 × 10⁵, ?4 × 10⁵ and ?6 × 10⁵ Pa at 2-day intervals. Plant height, leaf area and shoot dry weight of high and low nitrogen plants were reduced by lower osmotic potentials of the root medium. Osmotic stress caused greater shoot growth reduction in high N than in low N plants. Stressed and unstressed plants in 7.14 × 10⁴M N had more root dry matter than the corresponding plants in 28.6 × 10⁴M N. Dawn leaf water potential of stressed plants was 1 × 10⁵ to 5.5 × 10⁵ Pa lower than nutrient solution water potential. Nitrogen-deficient water-stressed plants, however, maintained higher dawn leaf water potential than high nitrogen water-stressed plants. It is suggested that this was due to higher root-to-shoot ratios of N deficient plants. The osmotic potentials of leaves at full turgor for control plants were about 1.3 × 10⁵ Pa higher in 7.14 × 10^?4M than in 28.6 × 10^?4M N and osmotic adjustment of 2.6 × 10⁵ and 4.3 × 10⁵ Pa was obtained in low and high N plants, respectively. The nitrogen status of plants, therefore, affected the ability of the rice plant to adjust osmotically during water stress. Plant water stress decreased transpiration and total N content in shoots of both N treatments. Reduced shoot growth as a result of water stress caused the decrease in amount of water transpired. Transpiration and N uptake were significantly correlated. Our results show that nitrogen content is reduced in water-stressed plants by the integrated effects of plant water stress per se on accumulation of dry matter and transpiring leaf area as well as the often cited changes in soil physical properties of a drying root medium.     

Evaluation of water stress control with polyethylene glycols by analysis of guttation

The water relations of pepper plants (Capsicum frutescens L.) under conditions conducive to guttation were studied to evaluate the control of plant water stress with polyethylene glycols. The addition of polyethylene glycol 6000 to the nutrient solution resulted in water relations similar to those expected in soil at the same water potentials. Specifically, xylem pressure potential in the root and leaf became more negative during a 24-hour treatment period, while osmotic potential of the root xylem sap remained constant. The decrease in pressure potential was closely correlated with the decrease in osmotic potential of the nutrient solution. In contrast, the addition of polyethylene glycol 400 to the nutrient medium resulted in a reduction of osmotic potential in the root xylem sap; this osmotic adjustment in the xylem was large enough to establish an osmotic gradient for entry of water and cause guttation at a nutrient solution osmotic potential of −4.8 bars. Pressure potential in the root and leaf xylem became negative only at nutrient solution osmotic potentials lower than −4.8 bars. About half of the xylem osmotic adjustment in the presence of polyethylene glycol 400 was caused by increased accumulation of K⁺, Na⁺, Ca²⁺, and Mg²⁺ in the root xylem. These studies indicate that larger polyethylene glycol molecules such as polyethylene glycol 6000 are more useful for simulating soil water stress than smaller molecules such as polyethylene glycol 400.     

Effects of repeated application of water stress on water status and growth of wheat

The effects on water status and growth of controlled cycles of water stress applied at various stages of development were studied on a semi-dwarf spring wheat (Triticum aestivum L.). The plants were grown in controlled environment chambers of the Duke University Phytetron at 24/18°C with a 12-h photo-period at about 600 μE m^?2 s^?1. Groups of plants were subjected to severe water stress by withholding irrigation, beginning at the 7th leaf, early anthesis, or early dough stages of development. A second cycle started 9 to 13 days after termination of the first cycle and maintained until the flag leaf water potential reached –25 bars at each of the growth stages. The lower leaves showed sign of wilting as indicated by curling in the first drying cycle at –7 bars and in the second cycle at –9 bars of leaf water potential during all stages of growth. Although these leaves recovered completely upon rewatering, onset of senescence was accelerated by three days in stressed plants. A preliminary drying cycle did not increase the ability of the plants to withstand subsequent stress because of severity of stress. Water stress of –25 bars at all three stages of growth reduced seed yield. The reduction was greater when a second stress cycle was also applied. Stress applied during early anthesis stage produced the smallest and the least number of seeds. The lack of osmotic adjustment probably was due to very rapid and severe development of water stress.     

Influence of Rate of Development of Leaf Water Deficits upon Photosynthesis, Leaf Conductance, Water Use Efficiency, and Osmotic Potential in Sorghum

This study reports the effect of rate of development of leaf water deficits in soil-grown sorghum (Sorghum bicolor) on the relationship of net photosynthesis, leaf conductance, and water use efficiency to leaf water potential, and on the degree of solute accumulation (osmotic adjustment). Recovery of these processes on rewatering, and responses during a second stress cycle were also studied. The most rapid rate of stress (1.2 MPa day^?1) resulted in no solute accumulation and the lowest rate of net photosynthesis and leaf conductance for any given leaf water potential during stress. Stress at 0.7 and 0.15 MPa day^?1 led to equal solute accumulations of approximately 0.6 MPa, but net photosynthesis, leaf conductance, and water use efficiency at a given leaf water potential were lower with the faster rate of stress (0.7 MPa day^?1). Additionally, leaf conductance at a given leaf turgor potential was lowest at the 1.2 MPa day^?1 stress rate, slightly higher at the intermediate rate of stress, and clearly highest at the slowest rate of stress. Recovery of both net photosynthesis and leaf conductance upon rewatering was rapid, taking less than 3 days, but full recovery of osmotic potential took between 6 and 11 days. One slow stress cycle had no influence on relationships during a second cycle. The concept of a threshold leaf water potential for stomatal closure is discussed and the conclusion reached that stomatal closure occurs slowly over a wide range of leaf water potential (> 1.0 MPa), the range being greater for slower rates of stress.     

Biomass production,photosynthesis, and leaf water relations of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> under moderate water stress

The perennial smooth cordgrass, Spartina alterniflora, has been successfully introduced in salty ecosystems for revegetation or agricultural use. However, it remains unclear whether it can be introduced in arid ecosystems. The aim of this study was to investigate the physiological response of this species to water deficiency in a climate-controlled greenhouse. The experiment consisted of two levels of irrigation modes, 100 and 50% field capacities (FC). Although growth, photosynthesis, and stomatal conductance of plants with 50% FC were reduced at 90 days from the start of the experiment, all of the plants survived. The water-stressed plants exhibited osmotic adjustment and an increase in the maximum elastic modulus that is assumed to be effective to enhance the driving force for water extraction from the soil with small leaf water loss. An increase in the water use efficiency was also found in the water-stressed plants, which could contribute to the maintenance of leaf water status under drought conditions. It can be concluded that S. alterniflora has the capacity to maintain leaf water status and thus survive in arid environment.     

An Association between Flowering and Reduced Stomatal Sensitivity to Water Stress in Pearl Millet [Pennisetum americanum (L.) Leeke]

Stomatal sensitivity to water stress was investigated in pearlmillet [Pennisetum americanum (L.) Leeke] in relation to stageof plant development, leaf water status and ABA content by samplingplants at midday. For the same leaf water potential (), droughtedplants with emerged panicles were found to have a greater leafconductance (g_L), indicative of greater stomatal opening, thanplants sampled prior to panicle emergence. The difference betweensuch flowering (F) and non-flowering (NF) plants in at stomatalclosure was estimated to be at least 0.6 MPa. This differencewas considered unlikely to be the result of differential bulkleaf osmotic adjustment, and for most samples from both F andNF plants, bulk leaf turgor potential (_p) was estimated to bezero. Stomatal closure in NF plants was associated in two genotypes(BJ 104 and line 112) with higher leaf ABA levels. Differencesin ABA levels between F and NF plants were, however, smalleror absent in genotypes Serere 39 and B282. These genotypes wereat lower than BJ 104 and line 112 when sampled and showed smallerdifferences between F and NF plants in conductance. Lower ABA levels in F plants are ascribed either to effectsof leaf ageing or to effects of flowering on ABA content ofthe leaf. Significant differences in g_L in the absence of differencesin ABA content are taken to imply changes in stomatal sensitivityto the hormone or in its access to the stomatal complex. Pennisetum americanum (L.) Leeke, pearl millet, flowering, stomata, water stress, abscisic acid     

Plant Growth in Polyethylene Glycol Solutions in Relation to the Osmotic Potential of the Root Medium and the Leaf Water Balance

The changes in leaf extension, plant dryweight, leaf area, netassimilation rate (E), relative growth-rate (R_W), and relativeleaf growth-rate (R_L), have been studied for four species grownfor 2 weeks in solutions of polyethylene glycol 4000 of controlledosmotic potentials. All aspects of growth were decreased bydecreasing the osmotic potential (_sol) of the root medium andthe leaf water potential (), and ceased when _/ was greater than— 10 bars in bean, cotton, maize. These plants were moresusceptible than ryegrass to water stress. Growth of bean stoppedat equal to about —6 bars, but even at —10 barsryegrass was capable of some growth. Slight decrease in fromthe values in the control plants decreased growth during thefirst week but partial recovery was apparent during the secondweek's growth in solution culture, when leaf extension, E, R_Land R_W increased in plants subjected to stress. Examinationof the water balance, water potential, osmotic potential andturgor of the leaf in relation to relative water content suggeststhat recovery was related to increased turgor and that the abilityof the plants to grow at reduced values of the osmotic potentialof the root medium and of the leaf water potential depend onthe maintenance of turgor.     

Effect of water stress on growth, osmotic adjustment, cell wall elasticity and water-use efficiency in Spartina alterniflora

In the northern spring–summer season of 2004–2005, vegetative propagated plants of Spartina alterniflora were grown under control and water stress conditions on the Mediterranean sea shore of the south-east of Tunis. Control plants were irrigated every week and water stress plants were irrigated until the soil achieved 50% (mild stress) and 25% (severe stress) field capacity (FC). Dry and fresh weight at the whole plant level (g plant⁻¹), shoot to root ratio on dry and fresh weight, photosynthesis (A), transpiration rate (E), instantaneous water-use efficiency (WUE_i), leaf water potential (Ψw), leaf water content (WC), osmotic potential at full turgor (Ψs¹⁰⁰), osmotic potential at turgor loss point (Ψs⁰), osmotic adjustment (OA), proline, sugars, inorganic compounds and cell wall elasticity (CWE) were evaluated during a period of 6 days period between 82 and 90 days after the beginning of treatment (DAT). Plants grown under severe and mild-water stress showed lower Ψw than in control plants with values that averaged −3.1, −1.6 and −0.9 MPa, respectively. S. alterniflora plants submitted to mild-water stress exhibited OA and a decrease in CWE. However, under severe water stress the OA was not observed and CWE also decreased, but it was higher than in the mild-water stress. OA was mainly explained by the accumulation of nitrates, sugars and at a lesser degree, proline. S. alterniflora had a strong decline of the dry and fresh weight of the whole plant associated to a marked decrease of photosynthesis (A) and transpiration (E) in response to water stress, although WUE_i was increased. These results suggest that OA and WUE_i can be important components of the water stress adaptation mechanism in this species, but they are not sufficient enough to contribute to resistance to water stress.     

Exogenous Ca<Superscript>2+</Superscript> alleviates nitrogen and water deficit,and improves growth of wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis>) seedlings exposed to high temperature

This study was designed to examine whether exogenous Ca²⁺ would improve nitrogen nutrition, water status and growth of high temperature (HT)-stressed wheat (Triticum aestivum) seedlings. Wheat plants were exposed to 35/30 and 25/20°C as temperature control. Some of HT-stressed plants were simultaneously treated with 4 mM Ca²⁺. External Ca²⁺ could obviously improve growth of HT-exposed wheat seedlings indicated by the biomass. Compared with Ca²⁺-untreated plants, total nitrogen content showed a significant increase in Ca²⁺-treated plants under HT stress, this primarily resulted from enhanced nitrate reductase activity and depressed loss of ammonium through photorespiration. External Ca²⁺ application could also increase leaf relative water content and alleviate osmotic stress via increased K⁺ ion and water-soluble carbohydrates in HT-stressed plants. Whereas free proline content showed remarkable decline in Ca²⁺-treated plants at HT stress.     

Effects of NaCl salinity and water stress on growth and leaf water relations of Asteriscus maritimus plants

Potted plants of Asteriscus maritimus (L.) Less were submitted to water stress (during two consecutive cycles, irrigation water was withheld for 5 days followed by a recovery period of 25 days) and saline stress (150 days of exposure to 0, 70 and 140 mM NaCl daily irrigation) in order to assess the effect on leaf water relations and growth parameters. Plants under saline and water stress conditions showed lower biomass and an early reduction in leaf expansion growth. Both stresses promoted a substantial degree of stomatal regulation; but, in spite of this, the plants showed signs of leaf tissue dehydration, decreases in RWC and Ψ_pd values. However, salt-treated plants, developed a NaCl inclusion mechanisms, underwent osmotic adjustment, which was able to maintain leaf turgor. Under both stress conditions g_l was independent to plant water status in the range between –0.8 and 1.0 MPa. Under water stress conditions, midday leaf water potential showed a threshold value (around −1.1 MPa), below which leaf conductance remained constant. In the salt-treated plants, the gradual closure of the stomata over a wide range of Ψ_md may be important in maintaining some level of photosynthesis.     

Leaf water characteristics and drought acclimation in sunflower genotypes

The responses of leaf water parameters to drought were examined using three sunflower (Helianthus annuus L.) genotypes. Osmotic potential at full water saturation (π¹⁰⁰), apoplastic water fraction (AWF) and bulk elastic modulus (BEM) were determined by pressure-volume curve analysis on well watered or on water-stressed plants (−1.0 MPa Ψ₁ < −1.5 MPa) previously drought-pretreated or not. The drought-pretreated plants were subjected to a 7-day drought period (predawn leaf water potential reached −0.9 MPa) followed by 8 days of rewatering. In well watered plants, all genotypes in response to drought acclimation displayed a significantly decreased π¹⁰⁰ associated with a decrease in the leaf water potential at the turgor-loss point (decrease in Ψ_tlp was between 0.15 and 0.21 MPa, depending on the genotype). In two genotypes, drought acclimation affected the partitioning of water between the apoplastic and symplastic fractions without any effect on the total amount of water in the leaves. As a third genotype displayed no modification of AWF and BEM after drought acclimation, the decreased π¹⁰⁰ was only due to the net accumulation of solutes and was consistent with the adjustment of the photochemical efficiency observed previously in this genotype in response to drought acclimation. In water-stressed plants, the osmotic adjustment (OA) can increase further beyond that observed in response to the drought pretreatment. However, the maintenance of photosynthetic rate and stomatal conductance at low leaf water potentials not only depends on the extent of osmotic adjustment, but also on the interaction between OA and AWF or BEM. Adaptative responses of leaf water parameters to drought are thus quite contrasted in sunflower genotypes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Leaf water relations characteristics of differently potassium fertilized and watered field grown barley plants

Seasonal leaf water relations characteristics were studied in fully irrigated spring barley (Hordeum distichum L. cv. Gunnar) fertilized at low (50 kg K ha⁻¹) or high (200 kg K ha⁻¹) levels of potassium applied as KCl. The investigation was undertaken from about 14 days before anthesis until the milk ripe stage in leaves of different position and age. Additionally, the effects of severe water stress on leaf water relations were studied in the middle of the grain filling period in spring barley (cv. Alis). The leaf water relations characteristics were determined by the pressure volume (PV) technique. Water relations of fully irrigated plants were compared in leaf No 7 with the water relations of slowly droughted plants (cv. Alis). Leaf osmotic potential at full turgor (ψ _π ¹⁰⁰ ) decreased 0.1 to 0.3 MPa in droughted leaves indicating a limited osmotic adjustment due to solute accumulation. The leaf osmotic potential at zero turgor (ψ _π ⁰ ) was about −2.2 MPa in fully irrigated plants and −2.6 MPa in droughted plants. The relative water content at zero turgor (R⁰) decreased 0.1 unit in severely droughted leaves. The ratio of turgid leaf weight to dry weight (TW/DW) tended to be increased by drought. The tissue modulus of elasticity (ε) decreased in droughted plants and together with osmotic adjustment mediated turgor maintenance during drought. A similar response to drought was found in low and high K plants except that the R⁰ and ε values tended to be higher in the high K plants. Conclusively, during drought limited osmotic adjustment and increase in elasticity of the leaf tissue mediated turgor maintenance. These effects were only slightly modified by high potassium application. The seasonal analysis in fully irrigated plants (cv. Gunnar) showed that within about 14 days from leaf emergence ψ _π ¹⁰⁰ decreased from about −0.9 to −1.6 MPa in leaf No 7 (counting the first leaf to emerge as number one) and from about −1.1 to −1.9 MPa in leaf No 8 (the flag leaf) due to solute accumulation. A similar decrease took place in ψ _π ⁰ except that the level of ψ _π ⁰ was displaced to a lower level of about 0.2 to 0.3 MPa. Both ψ _π ¹⁰⁰ and ψ _π ⁰ tended to be 0.05 to 0.10 MPa lower in high K than in low K plants. R⁰ was about 0.8 to 0.9 and was independent of leaf position and age, but tended to be highest in high K plants. The TW/DW ratio decreased from about 5.5 in leaf No 6 to 4.5 in leaf No 7 and 3.8 in leaf No 8. The TW/DW ratio was 4 to 10% higher in high K than in low K plants indicating larger leaf cell size in the former. The apoplastic water content (V_a) at full turgor constituted about 15% in leaf No 7. ε was maximum at full turgor and varied from about 11 to 34 MPa. ε tended to be higher in high K plants. Conclusively, in fully watered plants an ontogenetically determined accumulation of solutes (probably organic as discussed) occurred in the leaves independent of K application. The main effect of high K application on water relations was an increase in leaf water content and a slight decrease in leaf ψ_π. The effect of K status on growth and drought resistance is discussed.     

Salt tolerance of Beta macrocarpa is associated with efficient osmotic adjustment and increased apoplastic water content

The chenopod Beta macrocarpa Guss (wild Swiss chard) is known for its salt tolerance, but the mechanisms involved are still debated. In order to elucidate the processes involved, we grew wild Swiss chard exposed to three salinity levels (0, 100 and 200 mm NaCl) for 45 days, and determined several physiological parameters at the end of this time. All plants survived despite reductions in growth, photosynthesis and stomatal conductance in plants exposed to salinity (100 and 200 mm NaCl). As expected, the negative effects of salinity were more pronounced at 200 mm than at 100 mm NaCl: (i) leaf apoplastic water content was maintained or increased despite a significant reduction in leaf water potential, revealing the halophytic character of B. macrocarpa; (ii) osmotic adjustment occurred, which presumably enhanced the driving force for water extraction from soil, and avoided toxic build up of Na⁺ and Cl^– in the mesophyll apoplast of leaves. Osmotic adjustment mainly occurred through accumulation of inorganic ions and to a lesser extent soluble sugars; proline was not implicated in osmotic adjustment. Overall, two important mechanisms of salt tolerance in B. macrocarpa were identified: osmotic and apoplastic water adjustment.