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.     

Genotypic variation in tissue water relations of leaves and roots of black walnut (Juglans nigra) seedlings

Genetic variation in the drought response of leaf and root tissue water relations of seedlings of eight sources of black walnut ( Juglans nigra L.) was investigated using the pressure-volume technique. Tissue water relations were characterized at three stages of a drying cycle during which well-watered plants were allowed to desiccate and then were reirrigated.
Sources varied both in the capacity for and degree of leaf and root osmotic adjustment, and in the mechanism by which it was achieved. A decrease in osmotic potential at the turgor loss point (ψ_πp) of 0.4 MPa was attributable to increased leaf tissue elasticity in seedlings of four sources, while seedlings of an Ontario source exhibited a 0.7–0.8 MPa decline in ψ_πp as a result of both increased solute content and increased leaf tissue elasticity. Seedlings of a New York source showed no detectable osmotic adjustment.
In roots, decreased ψ_πp (osmotic potential at full hydration) and ψ_πp were observed under drought. Sources that exhibited significant leaf osmotic adjustment also generally showed a similar response in roots. Tissue elasticity and ψ_πp of roots were higher than those of shoots, whereas ψ_πp of the two organs was similar for most sources. Because of greater elasticity, roots exhibited a more gradual decline in turgor and total water potential than did leaves as tissue relative water content decreased.     

Salinity effects on the leaf water relations components and ion accumulation patterns in Avicennia germinans (L.) L. seedlings

Physiological traits involved in leaf water relations were evaluated in Avicennia germinans (L.) L. seedlings growing at different salinities in the field. Analysis of pressure-volume (P-V) curves and sap osmometry were combined to evaluate osmotic adjustment and cell elasticity, and the contribution of accumulated inorganic ions to osmotic potential was estimated. Seedlings growing in soils with interstitial water salinity above that of normal sea water showed a modification of the relationship between water potential and relative water content. Thus, their leaf osmotic potential at maximum turgor (Ψ_{π( max )}) and at zero turgor (Ψ_π(0)) was 1.41 and 1.82 MPa lower respectively, than that of the seedlings from the low salinity site. Volumetric moduli of elasticity () were between 17 and 23 MPa. Thus, ɛ was about 6 MPa lower in high-salinity plants indicating that their cells were slightly more elastic. Ionic concentration analysis showed that Σ [anions] and Σ [cations] were higher in the high-salinity site (22–35%) while the water content per unit dry mass was only 12–17% lower. Reduction in water content was insufficient to explain the increase in ion concentration. Ion concentration explained 73 and 66% of the osmotic potential estimated by P-V curves for leaves from low- and high-salinity sites, respectively. In conclusion, this study provided evidence that leaves of A. germinans seedlings adapt to hypersaline soils by increasing solute concentration by 52% and cell elasticity by 26%. Both processes allow leaf water uptake and turgor maintenance over a large range of soil water potential. Received: 30 June 1997 / Accepted 26 November 1997     

Osmotic adjustment inSorghum bicolor (L. Moench) grown under moisture stress in soil and osmotically modified solution cultures

Sorghum (Sorghum bicolor L. Moench) plants were grown in solution culture and stressed at three rates of decreasing leaf water potential (−0.123, −0.068 and −0.029 MPa day⁻¹) achieved by the incremental addition of an osmoticum, polyethylene glycol (PEG) 6000 to the solutions. Plants were also grown in soil and given different amounts of water which resulted in rates of decreasing leaf water potentials of −0.130 and −0.073 MPa day⁻¹. The rate of stress and the culture system influenced the accumulation of solutes in the cell, but not cell volume. A rapid stress (−0.123 and −0.130 MPa day⁻¹) to approximately −1.6 MPa leaf water potential resulted in 0.75 and 0.16 MPa of osmotic adjustment in the PEG and soil culture respectively. At moderate stress (−0.068 and −0.073 MPa day⁻¹) respective values were 1.68 and 0.58 MPa. There were some visual symptoms in the solution grown plants characteristic of uptake of high molecular weight PEG. However the relative growth rates of these plants were equal to or greater than those of the soil grown plants. In view of the differences in plant water status of soil and PEG solution cultured plants it was concluded that the use of the latter system would not be entirely suitable for some studies of drought resistance in sorghum, as related to crop performance in the field.     

The effect of potassium application on leaf water relations characteristics of field grown barley plants (Hordeum distichum L.)

Leaf water relations characteristics were studied in spring barley fertilized at low (50 kg ha^-1) or high (200 kg ha^-1) levels of potassium applied as KCl. The leaf water relations characteristics were determined by the pressure volume (PV) technique.Seasonal analysis in fully irrigated plants showed that within 2 weeks from leaf emergence the leaf osmotic potential at full turgor ( 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. ¹⁰⁰ was 0.05 to 0.10 MPa lower in high K than in low K plants. Thus, an ontogenetically determined accumulation of solutes occurred in the leaves independent of K application. The ratio of leaf weight at full turgor to dry weight (TW/DW) 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 tissue modulus of elasticity () was increased in high K plants. The main effect of high K application on water relations was an increase in leaf water content and a slight decrease in leaf 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.     

Sequence of drought response of maize seedlings in drying soil

Leaf elongation in monocotyledonous plants is sensitive to drought. To better understand the sequence of events in plants subjected to soil drying, leaf elongation and transpiration of maize seedlings ( Zea mays L.) of 4 cultivars were monitored continuously and the diurnal courses of the root and leaf water relations were determined. Results from this study indicate the following sequence of drought response: Leaf elongation decreased before changes in the leaf water relations of non‐growing zones of leaf blades were detected and before transpiration decreased. Reductions in leaf elongation preceded changes in the root water potential (ψ_w). Root ψ_w was not a very sensitive indicator of soil dryness, whereas the root osmotic potential (ψ_s) and root turgor (ψ_p) were more sensitive indicators. The earliest events observed in drying soil were a significant increase in the largest root diameter class (1 720 to 1 960 gm) and a decrease in leaf elongation ( P = 0.08) 2 days after withholding water. Significant increases in root length were observed 2 days later. Soil drying increased the number of fine roots with diameters of <240 µm. Slight increases in soil strength did not affect leaf elongation in the drying soil.     

Seasonal changes in production and nutrient content ofCynodon dactylon (L.) Pers. subjected to water deficits

Seasonal changes in water relations, production and mineral composition were studied in a sward ofCynodon dactylon (L). Pers. subjected to water deficits during a dry summer, and at recovery in autumn. The experiment was carried out under Mediterranean field conditions. Water deficits during summer reduced total dry matter production by 60%, but in autumn there were no differences between treatments. Compared to well-watered sward, the sward grown under drought showed an increase in potassium, calcium and nitrogen of 55, 10 and 10% respectively. These differences decreased with the arrival of autumn rains. Leaf osmotic potential (Ψ_Π) fell during the dry summer to −2.8 MPa in well-watered plants and to −4.2 MPa in stressed plants. In autumn there were no differences between treatments. Nevertheless, relative water content (RWC) only decreased to 0.86 in droughted plants. In summer potassium contributed to the osmotic adjustment. In contrast, under water deficits a decrease of 71% in sodium and, to a lesser but significant extent decreases in phosphorus, magnesium and chlorine was observed. Nitrogen, phosphorus and sulphur showed low concentrations during summer and increased in autumn.     

Leaf water relations characteristics of Lupinus angustifolius and L. cosentinii

Summary Lupins (Lupinus angustifolius and L. cosentinii) growing in 321 containers in a glasshouse were exposed to drought by withholding water. Leaf water potential (₁), and leaf osmotic potential (_s) were measured daily as soil water became depleted. Leaf water relations were further assessed by a pressure-volume technique and by measuring _s and relative water content of leaves after rehydration. Analysis by pressure-volume or cryoscopic techniques showed that leaf osmotic potential at saturation (_s100) decreased from -0.6 MPa in well watered to -0.9 MPa in severely droughted leaves, and leaf water potential at zero turgor (_zt) decreased from about -0.7 to -1.1 MPa in well watered and droughted plants, respectively. Relative water content at zero turgor (RWC_zt) was high (88%) and tended to be decreased by drought. The ratio of turgid leaf weight to dry weight was not influenced by drought and was high at about 8.0. The bulk elastic modulus () was approximately halved by drought when related to leaf turgor potential (_p) and probably mediated turgor maintenance during drought. The latter was found to be negatively influenced by rate of drought. Supplying the plants with high levels of K salts did not promote adjustment or turgor maintenance.     

Plant water relations as selection criteria for drought tolerance in mustard

Genotypes of mustard (B. juncea) were evaluated for concurrent changes in leaf water potential (Ψ), leaf osmotic potential (π), leaf turgor potential (P) and leaf relative water content (RWC) during moisture stress at reproductive stage of growth. The slope ‘b’ in the regression between Ψ and π varied from 0.43 to 0.97 and was positively correlated with P and RWC. The genotypes with ‘b’ around 0.7 were able to maintain P of about 0.5 MPa at Ψ of − 2.5 MPa and thus such value of ‘b’ seems to provide enough degree of tolerance against drought.     

Diurnal courses and factorial dependencies of leaf conductance and transpiration of differently potassium fertilized and watered field grown barley plants

During the grain filling period we followed diurnal courses in leaf water potential (ψ₁), leaf osmotic potential (ψ_π), transpiration (E), leaf conductance to water vapour transfer (g) and microclimatic parameters in field-grown spring barley (Hordeum distichum L. cv. Gunnar). The barley crop was grown on a coarse textured sandy soil at low (50 kg ha⁻¹) or high (200 kg ha⁻¹) levels of potassium applied as KCl. The investigation was undertaken at full irrigation or under drought. Drought was imposed at the beginning of the grain filling period. Leaf conductance and rate of transpiration were higher in the flag leaf than in the leaves of lower insertion. The rate of transpiration of the awns on a dry weight basis was of similar magnitude to that of the flag leaves. On clear days the rate of transpiration of fully watered barley plants was at a high level during most part of the day. The transpiration only decreased at low light intensities. The rate of transpiration was high despite leaf water potentials falling to rather low values due to high evaporative demands. In water stressed plants transpiration decreased and midday depression of transpiration occurred. Normally, daily accumulated transpirational water loss was lower in high K leaves than in low K leaves and generally the bulk water relations of the leaves were more favourable in high K plants than in low K plants. The factorial dependency of the flag leaf conductances on leaf water potential, light intensity, leaf temperature, and leaf-to-air water vapour concentration difference (ΔW) was analysed from a set of field data. From these data, similar sets of microclimatic conditions were classified, and dependencies of leaf conductance on the various environmental parameters were ascertained. The resulting mathematical functions were combined in an empirical simulation model. The results of the model were tested against other sets of measured data. Deviations between measured and predicted leaf conductance occurred at low light intensities. In the flag leaf, water potentials below-1.6 MPa reduced the stomatal apertures and determined the upper limit of leaf conductance. In leaves of lower insertion level conductances were reduced already at higher leaf water potentials. Leaf conductance was increased hyperbolically as photosynthetic active radiation (PAR) increased from darkness to full light. Leaf conductance as a function of leaf temperature followed an optimum curve which in the model was replaced by two linear regression lines intersecting at the optimum temperature of 23.4°C. Increasing leaf-to-air water vapour concentration difference caused a linear decrease in leaf conductance. Leaf conductances became slightly more reduced by lowered water potentials in the low K plants. Stomatal closure in response to a temperature change away from the optimum was more sensitive in high K plants, and also the decrease in leaf conductance under the influence of lowered ambient humidity proceeded with a higher sensitivity in high K plants. Thus, under conditions which favoured high conductances increase of evaporative demand caused an about 10% larger decrease in leaf conductance in the high K plants than in the low K plants. Stomatal sizes and density in the flag leaves differed between low and high K plants. In plants with partially open stomata, leaf conductance, calculated from stomatal pore dimensions, was up to 10% lower in the high K plants than in the low K plants. A similar reduction in leaf conductance in high K plants was measured porometrically. It was concluded that the beneficial effect of K supply on water use efficiency reported in former studies primarily resulted from altered stomatal sizes and densities.     

Moderate water stress affects tomato leaf water relations in dependence on the nitrogen supply

The responses of water relations, stomatal conductance (g_s) and growth parameters of tomato (Lycopersicon esculentum Mill. cv. Royesta) plants to nitrogen fertilisation and drought were studied. The plants were subjected to a long-term, moderate and progressive water stress by adding 80 % of the water evapotranspirated by the plant the preceding day. Well-watered plants received 100 % of the water evapotranspirated. Two weeks before starting the drought period, the plants were fertilised with Hoagland’s solution with 14, 60 and 110 mM NO₃ ⁻ (N14, N60 and N110, respectively). Plants of the N110 treatment had the highest leaf area. However, g_s was higher for N60 plants and lower for N110 plants. At the end of the drought period, N60 plants showed the lowest values of water potential (Ψ_w) and osmotic potential (Ψ_s), and the highest values of pressure potential (Ψ_p). N60 plants showed the highest Ψ_s at maximum Ψ_p and the highest bulk modulus of elasticity.     

Hydraulic resistance of two sorghums varying in drought resistance

Genotypes of sorghum [Sorghum bicolor (L.) Moench] vary in drought resistance. Yet it is not known if their hydraulic resistances vary. The objective of this study was to determine if the hydraulic resistance of a drought-resistant sorghum was the same as that of a drought-sensitive sorghum. Leaf water and osmotic potentials were measured daily, during a 14-d period, in leaves of a drought-resistant (‘KS9’) and a drought-sensitive (‘IA25’) sorghum, which had the roots in pots with a commercial potting soil that was either well watered or allowed to dry. Soil water potential, adaxial stomatal resistance, and transpiration rate were determined daily. Hydraulic resistance of the plants was calculated from the slope of the line relating soil water potential minus leaf water potential versus transpiration rate. When the soil was not watered, the drought-sensitive sorghum had a water potential that averaged −0.50 MPa lower and an osmotic potential that averaged −0.57 MPa lower, but a similar adaxial stomatal resistance (1.19 s mm⁻¹), compared with the drought-resistant sorghum. Seven days after the beginning of the experiment, the water potential of the soil with the drought-sensitive sorghum was −0.25 MPa lower than that of the soil with the drought-resistant sorghum. With the water-limited conditions, the drought-sensitive sorghum depleted the soil-water reserve more quickly and died 2 d before the drought-resistant sorghum. Under well watered conditions, the two sorghums had similar water potentials (−1.64 MPa), osmotic potentials (−2.83 MPa), and adaxial stomatal resistances (0.78 s mm⁻¹). The calculated hydraulic resistance of the two sorghums did not differ and averaged 3.4 × 10⁷ MPa s m⁻¹. The results suggested that the variation in susceptibility to drought between the two genotypes was due to differences in rate of soil-water extraction. Contribution No. 86-249-J from the Kansas Agricultural Experiment Station. The paper is dedicated to the memory of Dr Dan M Rodgers.     

Effects of water stress on the water relations of Phaseolus vulgaris and the drought resistant Phaseolus acutifolius

The tepary bean ( Phaseolus acutifolius Gray var. latifolius ), a drought resistant species, was compared under water stress conditions with the more drought susceptible P. vulgaris L. cvs Pinto and White Half Runner (WHR). In order to better understand the basis for the superior drought resistance of tepary, this study was designed to determine the relationships among leaf water potential, osmotic potential, turgor potential, and relative water content (RWC).
Plants were prestressed by withholding irrigation water. These stress pretreatments changed the relation between leaf water potential and relative water content of both species so that prestressed plants had lower water potentials than controls at the same leaf RWC. Tepary had lower water potentials at given RWC levels than Pinto or WHR; this can account for part of the superior resistance of tepary. In all genotypes, prestressed plants maintained osmotic potentials approximately 0.2 MPa lower than controls. Tepary reached osmotic potentials that were significantly lower (0.15 to 0.25 MPa) than Pinto or WHR. Both control and prestressed tepary plants had 0.05 to 0.25 MPa more turgor than Pinto or WHR at RWC values between 65 and 80%. Both prestressed and control tepary plants had greater elasticity (a lower elastic modulus) than Pinto or WHR. This greater turgor of tepary at low RWC values could be caused by several factors including greater tissue elasticity, active accumulation of solutes, or greater solute concentration.
Tepary had significantly lower osmotic potentials than the P. vulgaris cultivars, but there was little difference in osmotic potential between Pinto and WHR. Knowledge of differences in osmotic and turgor potentials among and within species could be useful in breeding for drought resistance in Phaseolus.     

Effect of nitrogen form on maize response to drought stress

Two hybrids of maize (Zea mays L.) differing in resistance to drought, were grown in chernozem soil in a greenhouse and were fertilized with two different forms of nitrogen: Ca(NO₃)₂ and (NH₄)₂SO₄ in concentrations corresponding to 100 kg of N ha^-1. After emergence of the 4th leaf, plants were exposed to drought. During the drought period, the parameters of plant water status (water potential, osmotic potential, turgor pressure and relative water content) and chlorophyll a+b concentration were monitored every two days. N and K concentration and accumulation over the drought period were also monitored.Next to differences in adaptability of the two hybrids to drought, the results demonstrate different adaptability of NH₄ and NO₃-treated plants within each hybrid. NH₄-plants of each hybrid maintain higher turgor pressure during the drought by better osmotic adaptation. Especially significant differences appear between chlorophyll (a+b) values of NH₄ and NO₃-treated plants and as affected by drought. Chlorophyll concentrations of NH₄-plants are higher than those of NO₃-plants both in control and droughted plants. NH₄ plants show a characteristic initial chlorophyll increase at the beginning of the drought period while in NO₃ plants chlorophyll constantly decreases throughout the whole drought period. The influence of the nitrogen form on chlorophyll concentration changes during drought does not appear to be affected by regulation of the K concentration.     

Effects of soil and air drought on growth,plant water status and leaf gas exchange in three Mediterranean cedar species: <Emphasis Type="Italic">Cedrus atlantica</Emphasis>, <Emphasis Type="Italic">C. brevifolia</Emphasis> and <Emphasis Type="Italic">C. libani</Emphasis>

Three- and four-year-old potted, greenhouse-grown cedar seedlings were subjected to two different watering regimes: half received full water supply and the other half was submitted to moderate drought (50% of the full water supply). Height growth was the greatest for C. atlantica and the most-limited for C. brevifolia in the well-watered set. However, in the dry set, height growth was less affected by drought conditions for C. brevifolia than for C. atlantica. Cedrus libani gave intermediate results for both watering regimes. Moderate drought provoked a decrease in osmotic potential at full leaf turgor and a long-lasting osmotic adjustment. When irrigation was withheld completely to induce severe soil drying, gas exchange decreased and then stopped at predawn water potentials of −3.0 MPa for C. brevifolia, between −2.6 and −2.8 MPa for C. libani, and at −2.4 MPa for C. atlantica, irrespective of watering regime. For all species, the dry set showed lower net photosynthesis (A) and stomatal conductance (g _s) than the plants in the well-watered set. A and g _s responded to variations in atmospheric water-vapour pressure deficit (VPD). As VPD increased, A and g _s decreased, and this trend was proportionate to initial values at low VPD, but remained independent of previous watering treatments, plant water status or species. To conclude, C. brevifolia appears to be a species with limited growth potential but strong soil drought tolerance whereas C. atlantica has strong growth potential when an adequate water supply is available but is more sensitive to soil drought. C. libani shows an intermediate behaviour for growth and drought tolerance.     

Water relations, gas exchange characteristics, and the level of some metabolites in two cultivars of spring wheat under different N regimes

Twenty eight-day old plants of two spring wheat cultivars differing in salinity tolerance were subjected to varying levels of nitrogen (56, 112, and 224 mg N·kg⁻¹ soil) for 42 days. Both cultivars performed differently under varying soil N levels in terms of growth, and grain yield and yield components. Nitrogen levels, 112 and 224 mg·kg⁻¹ soil, caused maximal growth in Sarsabz and Barani-83, respectively. Cv Sarsabz maintained higher leaf water and turgor potentials, but lower leaf osmotic potential than those of Barani-83 at all external N regimes. Sarsabz had higher Chl a, Chl b and carotenoids contents in leaves than those in Barani-83 at 56 and 112 mg N·kg⁻¹ soil. Sarsabz had higher contents of leaf soluble proteins, soluble sugars, and free amino acids than those in Barani-83 at all external N levels. In Barani-83 net CO₂ assimilation rate remained almost unchanged, whereas in Sarsabz it decreased consistently with increase in external N level. The better growth performance of Sarsabaz as compared to Barani-83 under varying soil N levels except 224 mg N·kg⁻¹ soil was associated with maintenance of high leaf turgor potential but not with net CO₂ assimilation rate.     

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.     

Day-night changes in leaf water relations associated with the rhythm of crassulacean acid metabolism in Kalanchoë daigremontiana

A study was made of the day-night changes under controlled environmental conditions in the bulk-leaf water relations of Kalanchoë daigremontiana, a plant showing Crassulacean acid metabolism. In addition to nocturnal stomatal opening and net CO₂ uptake, the leaves of well-watered plants showed high rates of gas exchange during the whole of the second part of the light period. Measurements with the pressure chamber showed that xylem tension increased during the night and then decreased towards a minimum at about midday; a significant increase in xylem tension was also seen in the late afternoon. Cell-sap osmotic pressure paralleled leaf malate content and was maximum at dawn and minimum at dusk. The relationship between these two variables indicated that the nocturnally synthesized malate was apparently behaving as an ideal osmoticum. To estimate bulk-leaf turgor pressure, values for water potential were derived by correcting the pressurechamber readings for the osmotic pressure of the xylem sap. This itself was found to depend on the malate content of the leaves. Bulk-leaf turgor pressure changed rhythmically during the day-night cycle; turgor was low during the late afternoon and for most of the night, but increased quickly to a maximum of 0.20 MPa around midday. In water-stressed plants, where net CO₂ uptake was restricted to the dark period, there was also an increase in bulk-leaf turgor pressure at the start of the light period, but of reduced magnitude. Such changes in turgor pressure are likely to be of considerable ecological importance for the water economy of crassulacean-acid-metabolism plants growing in their natural habitats.Abbreviation and symbols CAM Crassulacean acid metabolism - P turgor pressure - osmotic pressure - water potential Dedicated to Professor Dr. H. Ziegler on the occasion of his 60th birthday     

Seasonal Variations in Soil and Plant Water Status in a Quercus suber L. Stand: Roots as Determinants of Tree Productivity and Survival in the Mediterranean-type Ecosystem

Studies were conducted to examine changes in soil (Ψs) and plant water status during summer in a 16-year old Quercus suber plantation in southern Portugal. Continuous measurements were conducted between May 2003 and August 2004, while discontinuous measurements were conducted on a monthly basis between May and September 2003 and repeated between March and September 2004. Intensive measurements were conducted on five trees with mean height and DBH of 5.3 m and 11.6 cm, respectively, growing at close proximity to each other. Weather conditions and soil water potential (Ψs) at the rhizosphere of each of the trees measured at 0.3 and 1 m soil depth were continuously monitored. Predawn (Ψpd) and midday (Ψmd) leaf water potentials were determined every month. Soil and plant samples were also collected in June and September from different locations within the study site for δ¹⁸O isotope composition analysis. Pressure–volume (p–v) curves were constructed from plant shoots at different times during the vegetative period to determine osmotic potential at full saturation (Π¹⁰⁰), water potential at turgor loss point (Ψ_tlp), relative water content at turgor loss point (R*_tlp) and bulk modulus of elasticity (ε). Significant P < 0.05 decline in Ψs occurred between May and September, the lowest value recorded being –2.0 MPa. Decline in soil moisture affected tree water status, but decline in leaf water potential varied significantly (P < 0.05) among the trees. At the end of summer drought, lowest Ψpd measured was –1.7 MPa while the highest measured during this time was –0.8 MPa. Differences among trees were attributed to differences in rooting depth, as shown by regression analysis of ¹⁸O isotopes. Radial stem growth ceased when Ψs within the upper 0.3 m depth approached –1.5 MPa. The upper soil layers contributed approximately 33% of the total tree water requirement, between spring and mid summer when drought was experienced by trees. Deep soil layers however, supplied most of the water required during drought and no growth was recorded during this time. Stressed trees increased solute concentration of their tissues by a Magnitude of 0.7 MPa while bulk tissue elastic modulus increased by about 17 MPa. The study emphasizes the significance of roots as determinants of tree productivity and survival in the Mediterranean ecosystems.     

Drought Stress Response on Some Key Enzymes of Cowpea (<Emphasis Type="Italic">Vigna unguiculata</Emphasis> L. Walp.) Nodule Metabolism

A greenhouse experiment was carried out aiming to evaluate the response to drought stress of cowpea nodule enzymatic activities during different plant developmental stages leading to biological N₂ fixation. Stress was applied by controlling soil’s water-potential through a porous cup. Cowpea plants cv IPA 205 were grown in pots with yellow latosol soil under three different matric potential (ψ_m) treatments. Even with high evaporative demand and limited soil water availability, cowpea could not induce an extremely low leaf water potential (ψ_w). Sap ureides concentration in cowpea declined during the drought stress period. There was a decline in enzyme activity in the metabolic pathways concerned with N₂ fixation: NADH-dependent glutamate synthase (EC 1.4.1.14), glutamine synthetase (EC 6.3.1.2) and phosphoenolpyruvate carboxylase (EC 4.1.1.31). In contrast, an increase in glutamate dehydrogenase (EC 1.4.1.4) was observed as the ψ_m declined. Metabolism associated with N₂ assimilation was impaired every time that the ψ_w was reduced below −0.73 MPa as had happened in the stressed treatments. The stress applied by the porous cup was gradual and the plant recovered its turgor, avoiding permanent deleterious alterations in the cellular metabolism, even from a limited cowpea-growth ψ_m.