Effects of water stress cycles on turgor maintenance processes in pear leaves (Pyrus communis)

Effects of water deficits on leaf turgor maintenance processes were analyzed for pear trees (Pyrus communis L. cv. "Barlett") grown in 120-liter containers. Four irrigation treatments were applied: a well-watered control treatment, a spring water stress cycle (Sp), a summer water stress cycle (Su), and a spring plus summer water stress cycle (Sp + Su). For the Sp treatment, water application was progressively reduced from 100 to 20% of the control dose over a period of 27 days in spring. For the Su treatment, water application was progressively reduced over 23 days in summer, from 100 to 20% of the control dose. The Sp + Su treatment comprised both the spring and summer drought stress cycles. Pressure-volume (P-V) curves were constructed and stomatal conductances were determined for pear leaves from each treatment during the spring and summer stress cycles. Leaf water potential (Psi(pi) (0)) and relative water content (R(0)) at the turgor loss point of control leaves tended to decrease from spring to summer. Changes in leaf osmotic water potential at full turgor (Psi(pi) (100)) and in symplast water fraction (R(s)) did not explain the seasonal decrease in Psi(pi) (0). The water stress treatments had no effect on Psi(pi) (100), but R(s) was reduced by the water stress treatments, particularly during the summer stress cycle of the Su and Sp + Su treatments. The decrease in R(s) was correlated with an increase in the slope of the linear region of the P-V curve. Such a coupled adjustment would lead to increased water uptake capacity of water-stressed trees only under non-turgor conditions. Furthermore, pear leaves did not actively accumulate solutes. We conclude, therefore, that changes in leaf tissue water relations as a result of leaf acclimation to water stress are unlikely to facilitate maintenance of fruit productivity under drought.     

Clonal variation of Populus tremuloides responses to diurnal drought stress

We have developed an automated microprocessor controlled system for subjecting hydroponically grown plants to drought. Pumps and valves were used to move nutrient solutions into and out of a system of culture vessels in a growth chamber to provide periods of drought. Drought conditions were obtained by exposing the roots of hydroponically grown clones of aspen, Populus tremuloides Michx., to air in culture vessels temporarily emptied of nutrient medium. Over a 3-week period, the daily duration of drought was increased from 0 to 6 h. During this period, the plants became increasingly tolerant to drought, as shown by a decreasing propensity to wilt. All three clones sustained diurnal drought periods of 6 h for up to 5 weeks without detectable deterioration of health. Typical drought stress symptoms were observed including inhibition of growth, increased tissue amino acid content, and decreased water, solute, and turgor potentials in young leaves. In all clones, control plants had leaf water potentials between -1.0 and -1.6 MPa, whereas leaf water potentials of drought-treated plants were significantly lower, ranging from -1.7 to -3.0 MPa. Only one of the clones showed a significant decrease in leaf solute potential in response to drought. The decrease in leaf solute potential paralleled the decrease in water potential resulting in no significant difference in turgor potential. The other two clones had nonsignificant decreases to more negative leaf solute potentials under drought conditions resulting in significantly lowered turgor potentials. Leaf water potentials, solute potentials, and turgor potentials of the drought-treated plants returned to control values within two hours after rewatering. The growth inhibitions observed could not have been the consequence of loss of turgor. These results demonstrate genetic differences among aspen clones in water relations responses to drought.     

Physiological variation among western redcedar (Thuja plicata Donn ex D. Don) populations in response to short-term drought

? Variation in the ability of western redcedar (Thuja plicata Donn ex D. Don) populations to withstand water stress may exist because this species is found in coastal and interior biogeoclimatic subzones representing the full range of precipitation regimes in British Columbia, Canada.

? Seven western redcedar populations from locations in British Columbia, representing a wide range of habitat types, were assessed for their gas exchange and water relations response to controlled drought.

? Before drought, population variation occurred in stomatal conductance, net CO₂ assimilation rate and intrinsic water use efficiency and the relative water content at turgor loss point. During drought, populations had different responses of net CO₂ assimilation to decreasing predawn shoot water potential. After drought, populations differed in stomatal conductance and intrinsic water use efficiency, plus osmotic potential at turgor loss point, osmotic potential at saturation and apparent cuticular transpiration. Western redcedar populations from drier-inland habitats had a lower osmotic potential at turgor loss point, lower relative water content at turgor loss point and lower apparent cuticular transpiration in response to drought than populations from coastal origin with temperate maritime habitat.

? Reduction of cuticular water loss and adjustments of cellular water relations in response to drought was found to occur among seven western redcedar populations originating along a precipitation gradient while; there were minimal population differences in the gas exchange response to drought.

     

Impact of successive drought and re-watering cycles on growth and specific leaf area of two Populus x canadensis (Moench) clones, 'Dorskamp' and 'Luisa_Avanzo'

Responses to successive drought and re-watering cycles (1-3 cycles) were compared in greenhouse-grown cuttings of Populus x canadensis (Moench) clones, 'Luisa_ Avanzo' and 'Dorskamp.' Total leaf number increment rate, duration of leaf expansion, total and individual leaf area expansion rates and stomatal conductance were recorded periodically during the experiment. Soil water content (SWC) and predawn leaf water potential (Psi(WP)) were measured four times during each drought cycle. In parallel, relative leaf water content (RWC) and specific leaf area (SLA) were estimated on leaves collected from the top to bottom of each cutting. Under well-watered conditions, 'Luisa_Avanzo' and 'Dorskamp' differed in their patterns of leaf area expansion. Although duration of leaf expansion was similar between clones, 'Luisa_ Avanzo' exhibited higher total leaf number increment rates and individual leaf area increases than 'Dorskamp.' As a result, 'Luisa_Avanzo' cuttings reached larger individual and total leaf areas than 'Dorskamp.' 'Dorskamp' leaves had lower SLA than 'Luisa_Avanzo' leaves. In response to successive drought cycles, both clones underwent decreases in total leaf number increment rates and in total leaf area expansion rates, but both whole-plant and individual leaf areas were drastically reduced only in 'Luisa_Avanzo.' 'Dorskamp' maintained a constant leaf area as a result of an increase in the duration of leaf expansion during drought and a significant stimulation of individual leaf area expansion rate and total leaf number increment rate in response to re-watering. Drought caused a greater decrease in SLA in 'Luisa_Avanzo' than in 'Dorskamp.' Expanded leaves of 'Dorskamp' were constitutively dense or thick, or both, whereas leaves of 'Luisa_Avanzo' became dense or thick, or both, only in response to drought. In both clones, re-watering caused partial recovery of SLA to control values. Our data confirm previous field and greenhouse observations that 'Dorskamp' is more tolerant to successive drought and re-watering cycles than 'Luisa_Avanzo.' We conclude that this difference is mainly attributable to the plasticity of 'Dorskamp' after re-watering, rather than to clonal differences in drought responses.     

干旱胁迫对麻栎无性系蒸腾作用的影响

以13个3年生麻栎无性系为研究对象,采用盆栽试验方法,研究了在持续干旱胁迫条件下的麻栎无性系土壤水势、叶片水势光合速率和蒸腾速率变化规律及其关系。结果表明,随着干旱胁迫的加剧,13个麻栎无性系土壤水势呈现出不断下降的变化趋势,叶片水势的变化无一致规律,光合速率则先下降而后上升,蒸腾速率的变化规律不尽相同。在13个麻栎无性系中,2、4、5、7、9和16号无性系的蒸腾速率、叶片水势和光合速率具有相同规律,随着干旱胁迫的加剧先下降而后上升,具有较强的适应干旱逆境的能力。对不同干旱条件下无性系的蒸腾速率、土壤水势、叶片水势和光合速率的相关性分析发现,干旱胁迫7天时,蒸腾速率和光合速率达极显著正相关;干旱胁迫12天时,蒸腾速率和土壤水势间呈极显著负相关,光合速率与土壤水势呈显著负相关;干旱胁迫17天时,蒸腾速率与光合速率呈显著正相关。     

Physiological and morphological adaptations of the fruit tree Ziziphus rotundifolia in response to progressive drought stress

The physiological basis of drought resistance in Ziziphus rotundifolia Lamk., which is an important, multipurpose fruit tree of the northwest Indian arid zone, was investigated in a greenhouse experiment. Three irrigation regimes were imposed over a 34-day period: an irrigation treatment, a gradual drought stress treatment (50% of water supplied in the irrigation treatment) and a rapid drought stress treatment (no irrigation). Changes in gas exchange, water relations, carbon isotope composition and solute concentrations of leaves, stems and roots were determined. The differential rate of stress development in the two drought treatments did not result in markedly different physiological responses, but merely affected the time at which they were expressed. The initial response to decreasing soil water content was reduced stomatal conductance, effectively maintaining predawn leaf water potential (Psi(leaf)), controlling water loss and increasing intrinsic water-use efficiency, while optimizing carbon gain during drought. Carbon isotope composition (delta13C) of leaf tissue sap provided a more sensitive indicator of changes in short-term water-use efficiency than delta13C of bulk leaf tissue. As drought developed, osmotic potential at full turgor decreased and total solute concentrations increased in leaves, indicating osmotic adjustment. Decreases in leaf starch concentrations and concomitant increases in hexose sugars and sucrose suggested a shift in carbon partitioning in favor of soluble carbohydrates. In severely drought-stressed leaves, high leaf nitrate reductase activities were paralleled by increases in proline concentration, suggesting an osmoprotective role for proline. As water deficit increased, carbon was remobilized from leaves and preferentially redistributed to stems and roots, and leaves were shed, resulting in reduced whole-plant transpiration and enforced dormancy. Thus, Z. rotundifolia showed a range of responses to different drought intensities indicating a high degree of plasticity in response to water deficits.     

Genotypic variation in drought response of silver birch (Betula pendula): leaf water status and carbon gain

To assess genotypic variation in drought response of silver birch (Betula pendula Roth), we studied the plasticity of 16 physiological traits in response to a 12-14-week summer drought imposed on four clones in two consecutive years. In a common garden experiment, 1-year-old clonal trees from regions with low (550 mm year(-1)) to high rainfall (1270 mm year(-1)) were grown in 45-l pots, and leaf gas exchange parameters, leaf water potentials, leaf osmotic potentials and leaf carbon isotope signatures were repeatedly measured. There were no clonal differences in leaf water potential, but stomatal conductance (gs), net photosynthesis at ambient carbon dioxide concentration, photosynthetic water-use efficiency, leaf carbon isotope composition (delta13C) and leaf osmotic potentials at saturation (Pi0) and at incipient plasmolysis (Pip) were markedly influenced by genotype, especially gs and osmotic adjustment. Genotypes of low-rainfall origin displayed larger osmotic adjustment than genotypes of high-rainfall origin, although their Pi0 and Pip values were similar or higher with ample water supply. Genotypes of low-rainfall origin had higher gs than genotypes of high-rainfall origin under both ample and limited water supply, indicating a higher water consumption that might increase competitiveness in drought-prone habitats. Although most parameters tested were significantly influenced by genotype and treatment, the genotype x treatment interactions were not significant. The genotypes differed in plasticity of the tested parameters and in their apparent adaptation to drought; however, among genotypes, physiological plasticity and drought adaptation were not related to each other. Reduction of gs was the first and most plastic response to drought in all genotypes, and allowed the maintenance of high predawn leaf water potentials during the drought. None of the clones exhibited non-stomatal limitation of photosynthesis. Leaf gs, photosynthetic capacity, magnitude of osmotic adjustment and delta13C were all markedly lower in 2000 than in 1999, indicating root limitation in the containers in the second year.     

Seasonal variations in water relations in current-year leaves of evergreen trees with delayed greening

We investigated seasonal patterns of water relations in current-year leaves of three evergreen broad-leaved trees (Ilex pedunculosa Miq., Ligustrum japonicum Thunb., and Eurya japonica Thunb.) with delayed greening in a warm-temperate forest in Japan. We used the pressure-volume method to: (1) assess the extent to which seasonal variation in leaf water relations is attributable to leaf development processes in delayed greening leaves versus seasonal variation in environmental variables; and (2) investigate variation in leaf water relations during the transition from the sapling to the adult tree stage. Leaf mass per unit leaf area was generally lowest just after completion of leaf expansion in May (late spring), and increased gradually throughout the year. Osmotic potential at full turgor (Psi(o) (ft)) and leaf water potential at the turgor loss point (Psi(w) (tlp)) were highest in May, and lowest in midwinter in all species. In response to decreasing air temperature, Psi(o) (ft) dropped at the rate of 0.037 MPa degrees C(-1). Dry-mass-based water content of leaves and the symplastic water fraction of total leaf water content gradually decreased throughout the year in all species. These results indicate that reductions in the symplastic water fraction during leaf development contributed to the passive concentration of solutes in cells and the resulting drop in winter Psi(o) (ft). The ratio of solutes to water volume increased in winter in current-year leaves of L. japonicum and E. japonica, indicating that osmotic adjustment (active accumulation of solutes) also contributed to the drop in winter in Psi(o) (ft). Bulk modulus of elasticity in cell walls fluctuated seasonally, but no general trend was found across species. Over the growing season, Psi(o) (ft) and Psi(w) (tlp) were lower in adult trees than in saplings especially in the case of I. pedunculosa, suggesting that adult-tree leaves are more drought and cold tolerant than sapling leaves. The ontogenetic increase in the stress resistance of I. pedunculosa may be related to its characteristic life form because I. pedunculosa grows taller than the other species studied.     

Water and nutrient supply regimes and the water relations of juvenile leaves of Eucalyptus globulus

Pressure-volume analysis was used to study effects of irrigation and fertilization on the water relations of newly expanded juvenile leaves of Eucalyptus globulus Labill. seedlings growing in Portugal's Atlantic region. In May, at the beginning of the dry season, fertilization and irrigation treatments had no significant effects on the water relations parameters investigated. In September, at the end of the dry season, leaves from non-irrigated plants had a significantly higher apoplasmic water content and a higher dry weight/turgid weight ratio than leaves of similar physiological age from irrigated trees. The osmotic potential at full turgor and the water potential at the wilting point were lower in non-irrigated than in irrigated plants. Changes in osmotic potential at full turgor were negatively correlated with changes in dry weight/turgid weight ratio, suggesting that reductions in osmotic potential at full turgor were largely the result of decreases in cell size. Fertilization had no detectable effect on these variables or on leaf tissue bulk modulus of elasticity. Tissue elasticity was also unaffected by irrigation. Independently of water and nutrient supplies, leaf elasticity was higher and relative water content at the wilting point lower in leaves sampled in May than in leaves sampled in September. In non-irrigated plants, leaves sampled in September had a lower tissue elasticity and a lower osmotic potential at full turgor than leaves sampled in May, indicating that leaves produced at the end of the dry season generate lower water potentials as turgor is lost than leaves expanded early in the season.     

Leaf water relations and stomatal behavior of four allopatric Eucalyptus species planted in Mediterranean southwestern Australia

In 1986, four allopatric Eucalyptus species (E. camaldulensis Dehnh, E. saligna Smith, E. leucoxylon F. Muell and E. platypus Hook.) were planted together in a 480-mm rainfall zone, in 8-m wide contour belts as part of a plan to minimize waterlogging and secondary salinization. Throughout 1997, 1998 and 1999, there was significant inter-specific variation in predawn leaf water potential (Psi(pd)); however, maximum stomatal conductance (g(sm)) only differed significantly between species in mid to late summer. Relationships between g(sm) and Psi(pd) were significant and showed that stomata of E. camaldulensis were significantly more sensitive to Psi(pd), and presumably soil water potential, than stomata of E. leucoxylon or E. platypus. When applied to the Psi(pd) data, these relationships predicted that g(sm), and by inference transpiration, varied much less between species than Psi(pd). Diurnal measurements throughout the season confirmed this prediction, and showed that E. camaldulensis and E. saligna avoided drought by gaining access to deeper water, whereas E. leucoxylon and E. platypus maintained greater g(sm) at a given water stress than E. camaldulensis or E. saligna. Osmotic potentials measured after rehydration and water release curves of the leaves indicated that different mechanisms accounted for the apparent drought tolerance of E. leucoxylon and E. platypus. In summer, E. leucoxylon reduced osmotic potential at full and zero turgor by similar amounts compared with winter. In summer, E. platypus had a significantly lower bulk elastic modulus and relative water content at turgor loss point than E. camaldulensis, E. saligna or E. leucoxylon. This elastic adjustment resulted in a larger difference between osmotic potential at full and zero turgor in summer than in winter. The inherently low osmotic potential in E. leucoxylon and elastic adjustment in E. platypus resulted in turgor loss at a similar and significantly lower water potential than in E. camaldulensis or E. saligna. These results have implications for species selection for planting to manage groundwater recharge in areas prone to waterlogging and secondary salinization.     

Physiological traits of two Populus x euramericana clones,Luisa Avanzo and Dorskamp,during a water stress and re-watering cycle

We compared responses to drought and re-watering of greenhouse-grown cuttings of Populus x euramericana (Dode) Guinier clones, Luisa Avanzo and Dorskamp. Total leaf area, leaf number, leaf area increment and stomatal conductance were evaluated periodically during a 29-day drought period and for 16 days after re-watering. Soil water content and predawn leaf water potential (Psi(wp)) were measured on Days 29 and 45. On the same days, relative water content (RWC), specific leaf area (SLA), nitrogen, chlorophyll, soluble sugars, total phenols, flavanols and antioxidant activity were determined for leaves taken from the bottom to the top of each cutting. Leaves of Luisa Avanzo cuttings grew more rapidly than leaves of Dorskamp and exhibited higher SLA, but lower concentrations of nitrogen, chlorophyll and soluble sugars and lower antioxidant activity per unit area. On Day 29, after withholding water, both clones had closed their stomata, reduced rates of leaf growth, and lower Psi(wp) and RWC; however, the clones differed in their responses to soil water depletion. Compared to Dorskamp, Luisa Avanzo closed its stomata earlier and maintained higher Psi(wp), but lower RWC and leaf sugar concentrations. Antioxidant activity of leaf methanolic extracts decreased in response to water stress only in Luisa Avanzo. Leaf physiology and its modulation by water stress were age dependent in Luisa Avanzo.     

Water relations of seedlings of three Quercus species: variations across and within species grown in contrasting light and water regimes

We compared seedling water relations of three Mediterranean Quercus species (the evergreen shrub Q. coccifera L., the evergreen tree Q. ilex L. subsp. ballota (Desf.) Samp. and the deciduous or marcescent tree Q. faginea L.). We also explored seedling potential for acclimation to contrasting growing conditions. In March, 1-year-old seedlings of the three species were planted in pots and grown outdoors in a factorial combination of two irrigation regimes (daily (HW) and alternate day watering (LW)) and two irradiances (43 and 100% of full sunlight). At the end of July, predawn and midday water potentials (Psi(pd), Psi(md)) were measured, and pressure-volume (P-V) curves were obtained for mature current-year shoots. Species exhibited similar Psi(pd) and Psi(md) values, but differed in leaf morphology and water relations. The evergreens possessed larger leaf mass per area (LMA) and were able to maintain positive turgor pressure at lower water potentials than the deciduous species because of their lower osmotic potential at full turgor. However, the three species had similar relative water contents at the turgor loss point because Q. faginea compensated for its higher osmotic potential with greater cell wall elasticity. Values of Psi(pd) had a mean of -1.12 MPa in LW and -0.63 MPa in HW, and Psi(md) had a mean of -1.13 MPa in full sunlight and -1.64 MPa in shade, where seedlings exhibited lower LMA. However, the P-V curve traits were unaffected by the treatments. Our results suggest that Q. faginea seedlings combine the water-use characteristics of mesic deciduous oak and the drought-tolerance of xeric evergreen oak. The ability of Q. coccifera to colonize drier sites than Q. ilex was not a result of higher drought tolerance, but rather may be associated with other dehydration postponement mechanisms including drought-induced leaf shedding. The lack of treatment effects may reflect a relatively low contrast between treatment regimes, or a low inherent responsiveness of these traits in the study species, or both.     

Responses to water stress in two Eucalyptus globulus clones differing in drought tolerance

We evaluated drought resistance mechanisms in a drought-tolerant clone (CN5) and a drought-sensitive clone (ST51) of Eucalyptus globulus Labill. based on the responses to drought of some physiological, biophysical and morphological characteristics of container-grown plants, with particular emphasis on root growth and hydraulic properties. Water loss in excess of that supplied to the containers led to a general decrease in growth and significant reductions in leaf area ratio, specific leaf area and leaf-to-root area ratio. Root hydraulic conductance and leaf-specific hydraulic conductance decreased as water stress became more severe. During the experiment, the drought-resistant CN5 clone maintained higher leaf water status (higher predawn and midday leaf water potentials), sustained a higher growth rate (new leaf area expansion and root growth) and displayed greater carbon allocation to the root system and lower leaf-to-root area ratio than the drought-sensitive ST51 clone. Clone CN5 possessed higher stomatal conductances at moderate stress as well as higher hydraulic conductances than Clone ST51. Differences in the response to drought in root biomass, coupled with changes in hydraulic properties, accounted for the clonal differences in drought tolerance, allowing Clone CN5 to balance transpiration and water absorption during drought treatment and thereby prolong the period of active carbon assimilation.     

Reduction in turgid water volume in jack pine,white spruce and black spruce in response to drought and paclobutrazol

Significant reductions in needle water content were observed in white spruce (Picea glauca (Moench) Voss), black spruce (Picea mariana (Mill) B.S.P.), and jack pine (Pinus banksiana Lamb.) seedlings in response to a 10-day drought, although turgor was apparently maintained. When the seedlings were re-watered after the drought, jack pine needles regained their original saturated volume, whereas white spruce and black spruce needles did not. Significant drought-induced reductions in turgor-loss volume (i.e., tissue volume at the point of turgor loss) were observed in shoots of all three species, especially jack pine. Repeated exposure to 7 days of drought or treatment with the cytochrome P(450) inhibitor, paclobutrazol ((2RS,3RS)-1-(4-chlorophenyl)-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-pentan-3-ol), reduced seedling height relative to that of untreated controls in all three species. The reductions in saturated and turgor-loss needle volumes in the paclobutrazol-treated seedlings were comparable with those of seedlings subjected to a 10-day drought. The treatment-induced reductions in shoot and needle water contents enabled seedlings to maintain turgor with tissue volumes close to, or below, the turgor-loss volume of untreated seedlings. Paclobutrazol-treated seedlings subsequently survived drought treatments that were lethal to untreated seedlings.     

Functional and genetic characterization of gas exchange and intrinsic water use efficiency in a full-sib family of Pinus pinaster Ait. in response to drought

Drought is an important environmental factor in Mediterranean ecosystems affecting seedling recruitment, productivity or susceptibility to fires and pathogens. Studying water use efficiency in these environments is crucial due to its adaptive value allowing trees to cope with low water availability. We studied the phenotypic variability and genetic control of intrinsic water use efficiency (WUE(i)) and related traits in a full-sib family of Pinus pinaster under drought imposition. We detected significant differences in WUE(i) between clones of the same family and moderate heritability estimates that indicate some degree of genetic control over this trait. Stomatal conductance to water vapor was the trait most affected by drought imposition and it showed the strongest influence in WUE(i). Stomatal conductance to water vapor and specific leaf area (SLA) were the traits with highest heritabilities and they showed a significant genetic correlation with WUE(i), suggesting that selection of needles with low SLA values will improve WUE(i) in this species by reducing water losses through stomatal control.     

The impact of water and nutrient deficiencies on the growth, gas exchange and water relations of red oak and chestnut oak

Red oak (Quercus rubra), a mesic species, and chestnut oak (Quercus prinus), a xeric species, were grown in a greenhouse with and without fertilizer (F+ and F-, respectively) and subjected to a 10-week drydown (W-) or kept well watered (W+). In both species, fertilized seedlings exhibited greater reductions in mean net photosynthesis (A), leaf conductance (g(wv)), leaf water potential (Psi(leaf)) and water use efficiency (WUE) during the drydown than unfertilized seedlings. In the W- treatments, red oak showed greater reductions in A, g(wv) and Psi(leaf) than chestnut oak. Differential fertilization of the seedlings of both species had a greater effect on tissue water relations than differential watering. During the latter weeks of the drydown, there was no osmotic adjustment in red oak, but chestnut oak in the F+/W- treatment had significantly lower osmotic potentials at full and zero turgor than seedlings in any of the other treatments. The results indicate that high nutrient availability does not improve the drought tolerance of these two oak species.     

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

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

Drought adaptations and responses in five genotypes of Fraxinus pennsylvanica Marsh.: photosynthesis, water relations and leaf morphology

Genotypic variation in photosynthesis and plant water relations during drought, and in leaf and seedling morphology were examined in greenhouse-grown Fraxinus pennsylvanica Marsh. (green ash) from five populations located along an east-west transect from New York State to South Dakota. During a 17-day drought, South Dakota seedlings, from the most xeric habitat, maintained the highest net photosynthesis and leaf conductance, and New York seedlings, from the most mesic habitat, exhibited the lowest net photosynthesis and leaf conductance. All populations except New York adjusted osmotically during the 17-day drought, by the end of which New York seedlings had the highest osmotic potentials at full and zero turgor. Tissue elasticity increased in New York seedlings, but decreased in Nebraska seedlings during the drought. Leaves of South Dakota seedlings were the most xerophytic. They were smaller in area and greater in thickness and specific mass than leaves of other sources. Leaves of New York seedlings were thinner than those of the other genotypes and among the largest. Seedlings from South Dakota were smaller than those of the other populations.     

Comparative water relations of three successional hardwood species in central Wisconsin

Water relations of co-occurring understory saplings of Quercus ellipsoidalis E.J. Hill, an early successional, xeric species, Populus tremuloides Michx., an early successional, mesic species, and Acer rubrum L., a late successional species that occurs on both wet and dry sites, were evaluated on four dates during the 1986 growing season. The understory was characterized by high soil water content, low irradiance and low vapor pressure deficit throughout the growing season. Stomatal conductance and calculated transpiration flux were lowest for A. rubrum and highest for P. tremuloides and Q. ellipsoidalis. Except early in the growing season, leaf water potentials were lower in P. tremuloides than in the other species. Populus tremuloides had the highest bulk modulus of elasticity, Q. ellipsoidalis the lowest. Over the growing season, Populus tremuloides and Q. ellipsoidalis, but not A. rubrum, exhibited a decrease in osmotic potential at both full and zero turgor. Of the three species, Populus tremuloides exhibited the sharpest decrease in leaf water potential and turgor pressure with decreasing relative water content.     

Drought tolerance and transplanting performance of holm oak (Quercus ilex) seedlings after drought hardening in the nursery

Drought stress is the main cause of mortality of holm oak (Quercus ilex L.) seedlings in forest plantations. We therefore assessed if drought hardening, applied in the nursery at the end of the growing season, enhanced the drought tolerance and transplanting performance of holm oak seedlings. Seedlings were subjected to three drought hardening intensities (low, moderate and severe) for 2.5 and 3.5 months, and compared with control seedlings. At the end of the hardening period, water relations, gas exchange and morphological attributes were determined, and survival and growth under mesic and xeric transplanting conditions were assessed. Drought hardening increased drought tolerance primarily by affecting physiological traits, with no effect on shoot/root ratio or specific leaf mass. Drought hardening reduced osmotic potential at saturation and at the turgor loss point, stomatal conductance, residual transpiration (RT) and new root growth capacity (RGC), but enhanced cell membrane stability. Among treated seedlings, the largest response occurred in seedlings subjected to moderate hardening. Severe hardening reduced shoot soluble sugar concentration and increased shoot starch concentration. Increasing the duration of hardening had no effect on water relations but reduced shoot mineral and starch concentrations. Variation in cell membrane stability, RT and RGC were negatively related to osmotic adjustment. Despite differences in drought tolerance, no differences in mortality and relative growth rate were observed between hardening treatments when the seedlings were transplanted under either mesic or xeric conditions.