Leaf osmotic potential of Eucalyptus hybrids responds differently to freezing and drought, with little clonal variation

Concentrations of solutes, and thus leaf osmotic potential (Psi pi), often increase when plants are subject to drought or sub-zero (frost) temperatures. We measured Psi pi and concentrations of individual solutes in leaves of 3-year-old Eucalyptus camaldulensis Dehn., E. globulus Labill., E. grandis W. Hill ex Maid. and 29 hybrid clones on a site subjected to both summer drought and winter frost. We sought to characterize seasonal and genetic variations in Psi pi and to determine whether Psi pi or leaf turgor is related to bole volume increment. Leaf osmotic potential at full turgor (Psi pi(100)) was 0.7 MPa more negative in winter than in late summer, and this trend was uniform across genotypes. Soluble carbohydrates were confirmed as key contributors to Psi pi, accounting for 40-44% of total osmolality. The seasonal trend in Psi pi(100) was facilitated by changes in leaf morphology, such as reduced turgid mass:dry mass ratio and increased apoplastic water fraction in winter. Cell wall elasticity increased significantly from winter to summer. Our results suggest that elastic adjustment may be more important than osmotic adjustment in leaves exposed to drought. Although Psi pi(100) was a reasonable predictor of in situ osmotic potential and turgor, we found no relationship between any physiological trait and bole volume increment. Clone-within-family variation in Psi pi(100) was small in both summer and winter and was unrelated to bole volume increment. We conclude that, for the study species, tree improvement under water-limited conditions should concentrate on direct selection for growth rather than on indirect selection based on osmotic potential.     

Root temperature drives winter acclimation of shoot water relations in Cryptomeria japonica seedlings

In many temperate evergreen plant species, reductions in turgor loss point of leaves (Psi(tlp)) and leaf osmotic potential at full turgor (pi(sat)) occur from late summer to winter. To test the hypothesis that this seasonal change in leaf water relations is driven by root temperature, we manipulated the temperature of the roots and shoots of Cryptomeria japonica D. Don seedlings separately. Whole-plant warming diminished the seasonal changes in shoot water relations observed in the control plants, whereas shoot warming did not. Compared with the controls, root warming diminished the change in Psi(tlp) but not in pi(sat), whereas cooling accelerated the seasonal changes in shoot water relations. These results indicate that: (1) temperature responses of roots are involved in the seasonal changes in Psi(tlp) from late summer to winter; and (2) root temperature is partly responsible for the simultaneous changes in pi(sat). Whole-plant cooling caused increased root hydraulic resistance, suggesting that seasonal changes in shoot water relations represent adaptive responses to increased root hydraulic resistance at low root temperatures.     

Variations in transpiration rate and leaf cell turgor maintenance in saplings of deciduous broad-leaved tree species common in cool temperate forests in Japan

To clarify mechanisms underlying variation in transpiration rate among deciduous broad-leaved tree species, we measured diurnal changes in stomatal conductance (gs) and leaf water potential, and calculated the maximum transpiration rate (Emax), leaf-specific hydraulic conductance (K(s-l)) and difference between the soil water potential and the daily minimum leaf water potential (Psis - Psi(l,min)). Pressure-volume (P-V) measurements were made on leaves. Saplings of eight broad-leaved tree species that are common in Japanese cool temperate forests were studied. Maximum transpiration rate varied significantly among species. There was a statistically significant difference in Psis - Psi(l,min), but not in K(s-l). Species with large Emax also had large Psis - Psi(l,min) and gs. The results of the P-V analyses showed that species with a large Psis - Psi(l,min) maintained turgor even at low leaf water potentials. The similar daily minimum leaf pressure potentials (Psip) across all eight species indicate that Psip values below this minimum are critical. Based on these results, we suggest that the leaf cell capacity for turgor maintenance strongly affects Psis - Psi(l,min) and consequently Emax via stomatal regulation.     

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.     

Analysis of leaf water relations in leaves of two olive (Olea europaea) cultivars differing in tolerance to salinity

One-year-old rooted cuttings of olive (Olea europaea L. cvs. Frantoio and Leccino) were grown either hydroponically or in soil in a greenhouse. Plants were exposed to NaCl treatments (0, 100, and 200 mM) for 35 days, followed by 30 to 34 days of relief from salt stress to determine whether previously demonstrated genotypic differences in tolerance to salinity were related to water relations parameters. Exposure to high salt concentrations resulted in reductions in predawn water potential (Psi(w)), osmotic potential at full turgor (Psi(piFT)), osmotic potential at turgor loss point (Psi(piTLP)), and relative water content (RWC) in both cultivars, regardless of the growth substrate. Leaf Psi(w) and RWC returned to values similar to those of controls by the end of the relief period. The effect of salinity on Psi(pi) appeared earlier in Leccino than in Frantoio. Values for Psi(piFT) were -2.50, -2.87, and -3.16 MPa for the 0, 100, and 200 mM salt-treated Frantoio plants, respectively, and -2.23, -2.87, and -3.37 MPa for the corresponding Leccino plants. Recovery of Psi(pi) was complete for plants in the 100 mM salt treatment, but not for plants in the 200 mM salt treatment, which maintained an increased pressure potential (Psi(pi)) compared to control plants. Net solute accumulation was higher in Leccino, the salt-sensitive cultivar, than in Frantoio. In controls of both cultivars, cations contributed 39.9 to 42.0% of the total Psi(piFT), mannitol and glucose contributed 27.1 to 30.8%, and other soluble carbohydrates contributed 3.1 to 3.6%. The osmotic contribution of Na(+) increased from 0.1-2.1% for non-treated plants to 8.6-15.5% and 15.6-20.0% for the 100 mM and 200 mM salt-treated plants, respectively. The mannitol contribution to Psi(piFT) reached a maximum of 9.1% at the end of the salinization period. We conclude that differences between the two cultivars in leaf water relations reflect differences in the exclusion capacities for Na(+) and Cl(-) ions.     

Offsetting effects of reduced root hydraulic conductivity and osmotic adjustment following drought

Root hydraulic conductivity (L(p)) and leaf osmotic potential at full turgor (Psi(pi,o)) were measured in young, drought-stressed and nonstressed peach (Prunus persica (L.) Batsch), olive (Olea europaea L.), citrumelo (Poncirus trifoliata Raf. x Citrus paradisi Macf.) and pistachio (Pistachia integerrima L.). Drought stress caused a 2.5- to 4.2-fold reduction in L(p), depending on species, but Psi(pi,o) was reduced only in citrumelo and olive leaves by 0.34 and 1.4 MPa, respectively. No differences existed in L(p) among species for nonstressed plants. A simple model linking L(p) to osmotic adjustment through leaf water potential (Psi) quantified the offsetting effects of reduced L(p) and osmotic adjustment on the hypothetical turgor pressure difference between drought-stressed and nonstressed plants (DeltaPsi(p)). For olive, the 2.5-fold reduction in L(p) caused a linear decrease in DeltaPsi(p) such that the effect of osmotic adjustment was totally negated at Psi = -3.2 MPa. Thus, no stomatal closure would be required to maintain higher turgor in drought-stressed olive plants than in nonstressed plants over their typical diurnal range of Psi (-0.6 to -2.0 MPa). For citrumelo, osmotic adjustment was offset by reduced L(p) at Psi approximately -0.9 MPa. Unlike olive, stomatal closure would be necessary to maintain higher turgor in drought-stressed citrumelo plants than in nonstressed plants over their typical diurnal range of Psi (0 to -1.5 MPa). Regardless of species or the magnitude of osmotic adjustment, my analysis suggests that a drought-induced reduction in L(p) reduces or eliminates turgor maintenance through osmotic adjustment.     

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.     

Changes in the water relations of balsam fir and white birch saplings after thinning

In the autumn of 1987, young balsam fir (Abies balsamea (L.) Mill.) and white birch (Betula papyrifera Marsh.) trees were thinned and their water relations followed during the next two growing seasons. At the beginning of the first summer following treatment, thinned trees of both species had lower osmotic potentials at full saturation (Psi(pi,sat)) and at turgor loss point (Psi(pi,tlp)) compared with controls. At this time, Psi(pi,sat) was linearly related to the percentage of full sunlight reaching the trees. A higher sugar concentration in leaves was an important component of the lower Psi(pi,sat) of thinned trees. For the other two sampling dates during the first growing season after treatment and all three sampling dates during the second growing season after treatment, little osmotic adjustment of the thinned trees relative to the control tress was observed in either species. The absence of osmotic adjustment during the second growing season following thinning suggests that other mechanisms were responsible for the acclimation of the treated trees to the higher atmospheric evaporative demand. Sapwood permeability (k) of white birch was higher than that of balsam fir, but no differences in k or in sapwood area were found between treated and control trees of either species. Predawn water potentials (Psi(pred)) of treated trees were less negative than those of controls.     

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.     

Biophysical constraints on leaf expansion in a tall conifer

The physiological mechanisms responsible for reduced extension growth as trees increase in height remain elusive. We evaluated biophysical constraints on leaf expansion in old-growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees. Needle elongation rates, plastic and elastic extensibility, bulk leaf water (Psi(L)) and osmotic (Psi(pi)) potential, bulk tissue yield threshold and final needle length were characterized along a height gradient in crowns of > 50-m-tall trees during the period between bud break and full expansion (May to June). Although needle length decreased with increasing height, there was no height-related trend in leaf plastic extensibility, which was highest immediately after bud break (2.9%) and declined rapidly to a stable minimum value (0.3%) over a 3-week period during which leaf expansion was completed. There was a significant positive linear relationship between needle elongation rates and plastic extensibility. Yield thresholds were consistently lower at the upper and middle crown sampling heights. The mean yield threshold across all sampling heights was 0.12 +/- 0.03 MPa on June 8, rising to 0.34 +/- 0.03 MPa on June 15 and 0.45 +/- 0.05 MPa on June 24. Bulk leaf Psi(pi) decreased linearly with increasing height at a rate of 0.004 MPa m(-1) during the period of most rapid needle elongation, but the vertical osmotic gradient was not sufficient to fully compensate for the 0.015 MPa m(-1) vertical gradient in Psi(L), implying that bulk leaf turgor declined at a rate of about 0.011 MPa m(-1) increase in height. Although height-dependent reductions in turgor appeared to constrain leaf expansion, it is possible that the impact of reduced turgor was mitigated by delayed phenological development with increasing height, which resulted in an increase with height in the temperature during leaf expansion.     

Variation in growth and osmotic regulation of roots of water-stressed maritime pine (Pinus pinaster Ait.) provenances

Hydroponically cultivated Pinus pinaster Ait. seedlings of a drought-sensitive population from France (Landes) and of a more drought-adapted population from Morocco (Tamjoute) were subjected to a progressive increase in water stress by additions of an osmoticum (polyethylene glycol 600) to the nutrient solution. The final osmotic potential (Psi(ms)) of the nutrient solution was achieved over a period of up to 6 days, and ranged from -0.03 (control, no added osmoticum) to -0.8 MPa. In the 6 days during which water stress was imposed, roots elongated faster in the Moroccan provenance than in the French provenance, but the applied water deficits did not inhibit root elongation in either population. Among treatments, root dry weight per unit root length, total root dry weight and root/shoot dry weight ratio increased with decreasing Psi(ms) in both provenances. Both the water potential (Psi(w)) of the roots (apices) and the water potential difference between the roots and the nutrient solution decreased as Psi(ms) decreased. The reduction in Psi(w) was matched by a decrease of comparable magnitude in cell osmotic potential (Psi(pi)) so that root turgor was unaffected by the Psi(ms). Osmotic adjustment was greater, however, in the Moroccan provenance than in the French provenance. Consequently, under the osmotically imposed water stress, the water potential difference between root and nutrient solution was greater in the Moroccan provenance than in the French provenance. Similar changes in plant water relations were observed when seedlings were grown in drying sand.     

Physiological responses of radiata pine roots to soil strength and soil water deficit

We investigated physiological responses of radiata pine (Pinus radiata D. Don) roots to soil strength and soil water deficit by measuring the osmotic potential (Psi(pi)) and yield turgor (Y) in the elongation zone of root segments of seedlings growing (i) in polyethylene glycol 4000-containing rooting solution of different water potentials (Psi(s)) and (ii) in soil of different soil strengths (Q) at the same soil matric potential (Psi(m)). Root elongation rate (Deltal/Deltat) decreased progressively with decreasing Psi(s) and was associated with decreased Psi(pi) and decreased turgor pressure (P). Osmotic adjustment occurred at Psi(s) < -0.2 MPa. Over a range in Psi(s) of -0.01 to -1.0 MPa, Psi(pi) fell 0.3 MPa whereas P fell 0.7 MPa. Mean Psi in the solution experiment was 0.37 MPa and did not differ significantly with Psi(s) (P = 0.10). Root elongation rate decreased exponentially as Q increased from 0 to 3.0 MPa, and was associated with an increase in P of 0.11 MPa as a consequence of Psi(pi) decreasing by the same amount. Mean Y in the soil experiment was 0.49 MPa and did not change significantly with Q (P = 0.87).     

Shoot water relations of mature black spruce families displaying a genotype x environment interaction in growth rate. II. Temporal trends and response to varying soil water conditions

Pressure-volume curves and shoot water potentials were determined for black spruce (Picea mariana (Mill.) BSP) trees from four full-sib families at the Petawawa Research Forest, Ontario, Canada. Trees were sampled from a dry site in 1992 and from the dry site and a wet site in 1993. Modulus of elasticity (epsilon), osmotic potential at turgor loss point (Psi(tlp)) and relative water at turgor loss point (RWC(tlp)) all decreased during the growing season. Osmotic potential at saturation (Psi(sat)) and turgor displayed no general temporal trend. Across a range of environmental conditions, Female 59 progeny had equal or lower Psi(sat), and higher or similar epsilon, mean turgor pressure (P(x)) and predawn turgor pressure (P(pd)) compared with Female 63 progeny. Osmotic potential at saturation decreased as water stress increased from mild to moderate and increased as water stress increased from moderate to severe. Stable genetic differences in Psi(sat) were maintained by the same rate of osmotic adjustment from low to moderate water stress. Modulus of elasticity and RWC(tlp) decreased with decreasing water availability, whereas Psi(tlp) showed no response. The combined effects of Psi(sat) and epsilon resulted in no change in P(pd) as water stress increased from low to moderate values, but turgor declined sharply as water stress increased from moderate to high values. We conclude that drought tolerance traits strongly influence the growth of these black spruce families across sites of varying water availability.     

Gas exchange, water relations and osmotic adjustment in Phillyrea latifolia grown at various salinity concentrations

Leaf gas exchange, water relations and osmotic adjustment were studied in hydroponically grown Phillyrea latifolia L. plants exposed to 5 weeks of salinity stress (0, 80, 160, 240 and 320 mM NaCl) followed by 5 weeks of treatment with half-strength Hoagland solution. Whole-plant relative growth rate and root/shoot and lateral/structural root ratios were also evaluated. Net CO2 assimilation rate, stomatal conductance and transpiration rate were markedly decreased by all of the salt treatments. Growth was also strongly depressed by all salt treatments, especially lateral root growth. Leaf water potential decreased soon after salinity stress was imposed, whereas there was a lag of several weeks before leaf osmotic potential decreased in response to the salt treatments. After 5 weeks of salinization, leaf turgor of salt-treated plants was similar to that of controls. Although Na+ + Cl- contributed little to the salt-induced changes in osmotic potential at full turgor (Psi(piFT)), the contributions of K+, mannitol (Man) and glucose (Glc) to Psi(piFT) markedly increased as external salinity increased. Salt accumulation was negligible in the youngest leaves, which mostly accumulated soluble carbohydrates and K+; in contrast, old leaves served as storage sinks for Na+ and Cl-. Photosynthetic performance of salt-treated plants fully recovered once salt was leached from the root zone, with the recovery rate depending on the severity of the salt stress previously experienced by the plants. Recovery of gas exchange occurred even though the leaves still had a salt load similar to that detected in leaves at the end of the 5-week salinity period, and had markedly lower concentrations of K+ and soluble carbohydrates than control leaves. We conclude that salt-induced water stress primarily controlled gas exchange of salt-treated P. latifolia leaves, whereas the salt load in the leaves did not cause irreversible damage to the photosynthetic apparatus.     

Factors involved in alleviating water stress by partial crop removal in pear trees

We studied the relief of water stress associated with fruit thinning in pear (Pyrus communis L.) trees during drought to determine what mechanisms, other than stomatal adjustment, were involved. Combinations of control irrigation (equal to crop water use less effective rainfall) and deficit irrigation (equal to 20% of control irrigation), fruit load (unthinned and thinned to 40 fruits per tree) and root pruning (pruned and unpruned) treatments were applied to pear (cv. 'Conference') trees during Stage II of fruit development. Daily patterns of midday stem water potential (Psi(stem)) and leaf conductance to water vapor (g(l)) of deficit-irrigated trees differed after fruit thinning. In response to fruit thinning, gl progressively declined with water stress until 30 days after fruit thinning and then leveled off, whereas the effects of decreased fruit load on Psi(stem) peaked 30-40 days after fruit thinning and then tended to decline. Soil water depletion was significantly correlated with fruit load during drought. Our results indicate that stomatal adjustment and the resulting soil water conservation were the factors determining the Psi(stem) response to fruit thinning. However, these factors could not explain differences in daily patterns between g(l) and Psi(stem) after fruit thinning. In all cases, effects of root pruning treatments on Psi(stem) in deficit-irrigated trees were transitory (Psi(stem) recovered from root pruning in less than 30 days), but the recovery of Psi(stem) after root pruning was faster in trees with low fruit loads. This behavior is compatible with the concept that the water balance (reflected by Psi(stem) values) was better in trees with low fruit loads compared with unthinned trees, perhaps because more carbon was available for root growth. Thus, a root growth component is hypothesized as a mechanism to explain the bimodal Psi(stem) response to fruit thinning during drought.     

Response of five temperate deciduous tree species to water stress

Gas exchange, tissue water relations, and leaf/root dry weight ratios were compared among young, container-grown plants of five temperate-zone, deciduous tree species (Acer negundo L., Betula papyrifera Marsh, Malus baccata Borkh, Robinia pseudoacacia L., and Ulmus parvifolia Jacq.) under well-watered and water-stressed conditions. There was a small decrease (mean reduction of 0.22 MPa across species) in the water potential at which turgor was lost (Psi(tlp)) in response to water stress. The Psi(tlp) for water-stressed plants was -1.18, -1.34, -1.61, -1.70, and -2.12 MPa for B. papyrifera, A. negundo, U. parvifolia, R. pseudoacacia, and M. baccata, respectively. Variation in Psi(tlp) resulted primarily from differences in tissue osmotic potential and not tissue elasticity. Rates of net photosynthesis declined in response to water stress. However, despite differences in Psi(tlp), there were no differences in net photosynthesis among water-stressed plants under the conditions of water stress imposed. In A. negundo and M. baccata, water use efficiency (net photosynthesis/transpiration) increased significantly in response to water stress. Comparisons among water-stressed plants showed that water use efficiency for M. baccata was greater than for B. papyrifera or U. parvifolia. There were no significant differences in water use efficiency among B. papyrifera, U. parvifolia, A. negundo, and R. pseudoacacia. Under water-stressed conditions, leaf/root dry weight ratios (an index of transpiration to absorptive capacity) ranged from 0.77 in R. pseudoacacia to 1.05 in B. papyrifera.     

Water relations of several hardwood species in response to throughfall manipulation in an upland oak forest during a wet year

We investigated the effects of altered precipitation on leaf osmotic potential at full turgor (Psi(pio)) of several species in an upland oak forest during the 1994 growing season as part of a Throughfall Displacement Experiment at the Walker Branch Watershed near Oak Ridge, Tennessee. The main species sampled included overstory chestnut oak (Quercus prinus L.), white oak (Q. alba L.), red maple (Acer rubrum L.); intermediates sugar maple (A. saccharum L.) and blackgum (Nyssa sylvatica Marsh.); and understory dogwood (Cornus florida L.) and red maple. The precipitation treatments were: ambient precipitation; ambient minus 33% of throughfall (dry); and ambient plus 33% of throughfall (wet). Except in late September, midday leaf water potentials (Psi(l)) were generally high in all species in all treatments, ranging from -0.31 to -1.34 MPa for C. florida, -0.58 to -1.51 MPa for A. rubrum, and -0.78 to -1.86 MPa for Q. prinus. Both treatment and species differences in Psi(pio) were evident, with oak species generally exhibiting lower Psi(pio) than A. saccharum, A. rubrum, C. florida, and N. sylvatica. The Psi(pio) of C. florida saplings declined in the dry treatment, and Q. prinus, Q. alba, and A. saccharum all exhibited a declining trend of Psi(pio) in the dry treatment, although Psi(pio) of Q. prinus leaves increased in late August, corresponding to a recovery in soil water potential. Cornus florida exhibited osmotic adjustment with the largest adjustment coinciding with the period of lowest soil water potential in June. The only other species to exhibit osmotic adjustment was Q. prinus, which also maintained a lower baseline Psi(pio) than the other species. We conclude that a 33% reduction of throughfall is sufficient both to alter the water relations of some species in the upland oak forest and to enable the identification of those species capable of osmotic adjustment to a short-term drought during a wet year.     

Planting stress in newly planted jack pine and white spruce. 2. Changes in tissue water potential components

Water relations of bare-root jack pine (Pinus banksiana Lamb.) and white spruce (Picea glauca (Moench) Voss) planted in a greenhouse and on a boreal cut-over site were examined during the first growing season. In field-planted trees, maximum stomatal conductances (g(wv)) were initially low (< 0.10 cm s(-1)). Base and minimum xylem pressure potentials (Psi(x(base)) and Psi(x(min))) were less than -1.5 and -1.7 MPa for jack pine and -2.0 and -2.6 MPa for white spruce, respectively. During the growing season, maximum g(wv) increased in both species to around 0.2 cm s(-1). Base and minimum xylem pressure potentials also increased in both species to around -0.5 and -1.0 MPa in jack pine and -1.0 and -1.5 MPa in white spruce, respectively. Minimum xylem pressure potentials in white spruce fell below the turgor loss point during the first half of the growing season. Osmotic potential at the turgor loss point Psi(pi(TLP)) decreased after field planting to around -2.7 and -2.3 MPa in jack pine and white spruce, respectively. In the greenhouse, minimum values of Psi(pi(TLP)) were -2.2 and -2.3 MPa in jack pine and white spruce, respectively. Maximum bulk modulus of elasticity was greater in white spruce and underwent greater seasonal change than in jack pine. Relative water content (RWC) at turgor loss ranged between 71 and 74% in jack pine and 80 and 87% in white spruce. Available turgor (T(avail)), defined as the integral of turgor over the range of RWC between Psi(x(base)) and xylem pressure potential at the turgor loss point, was similar in jack pine and white spruce just after field planting. For the rest of the growing season, however, T(avail) in jack pine was two to three times that in white spruce. Diurnal turgor (T(diurnal)), defined as the integral of turgor over the range of RWC between Psi(x(base)) and Psi(x(min)), as a percent of T(avail) was higher in field-planted white spruce than jack pine until the end of the season. Dynamics of tissue water potential components are discussed in relation to plantation establishment.     

Growth, leaf morphology, water use and tissue water relations of Eucalyptus globulus clones in response to water deficit

Changes in leaf size, specific leaf area (SLA), transpiration and tissue water relations were studied in leaves of rooted cuttings of selected clones of Eucalyptus globulus Labill. subjected to well-watered or drought conditions in a greenhouse. Significant differences between clones were found in leaf expansion and transpiration. There was a significant clone x treatment interaction on SLA. Water stress significantly reduced osmotic potential at the turgor loss point (Pi0) and at full turgor (Pi100), and significantly increased relative water content at the turgor loss point and maximum bulk elastic modulus. Differences in tissue water relations between clones were significant only in the mild drought treatment. Among clones in the drought treatments, the highest leaf expansion and the highest increase in transpiration during the experiment were measured in those clones that showed an early and large decrease in Pi0 and Pi100.     

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.