Diurnal variation in leaf extension of Salix viminalis at two nitrogen supply rates

To investigate how nitrogen supply might affect the biophysical factors controlling diurnal variation in leaf extension, pot-grown Salix viminalis L. were supplied with nitrogen at a low relative addition rate of 0.05 g N g(-1) N day(-1) (low N) or were given free access to all nutrients (high N). Leaf extension, turgor pressure, turgor after stress relaxation and the plastic extensibility of leaf tissue were determined for growing leaves every 4 h during two days of clear skies in August. Plants in the high-N treatment had a significantly higher relative growth rate, dry weight, shoot/root ratio, leaf nitrogen concentration, total leaf area, final area of single leaves and epidermal cell size than plants in the low-N treatment. The periodicity of leaf extension was similar in both treatments with high values during the afternoon and early evening, and negligible values during the night and in the early morning. The maximum rate of leaf extension was higher in high-N than in low-N plants. Leaf water potential and leaf osmotic potential decreased in the morning and increased in the afternoon with highest values during the night. Calculated values of turgor pressure showed no consistent diurnal trend and did not correlate with the rate of leaf extension. There was no consistent difference in turgor between treatments. Turgor after stress relaxation varied diurnally. The difference between turgors before and after stress relaxation also varied diurnally and was largely in phase with the diurnal pattern of leaf extension. These data are consistent with either a causal role for growth turgor (difference between turgors before and after stress relaxation) in the regulation of cell expansion, or a diurnal variation in turgors after relaxation, attributable to different capacities for cell wall loosening at different times of day. Plastic extensibility of leaf tissue showed no diurnal pattern but consistently higher values were found in high-N than in low-N plants. We conclude that the effects of nitrogen supply on leaf water relations did not limit leaf extension, but that nitrogen supply did affect processes associated with cell wall loosening and enlargement. Nitrogen supply did not affect final values of turgor after relaxation, but it presumably affected the rate at which relaxation proceeded.     

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.     

Leaf growth, photosynthesis and tissue water relations of greenhouse-grown Eucalyptus marginata seedlings in response to water deficits

Leaf growth, rate of leaf photosynthesis and tissue water relations of shoots of Eucalyptus marginata Donn ex Sm. (jarrah) seedlings were studied during a soil drying and rewatering cycle in a greenhouse experiment. Rates of leaf growth and photosynthesis were sensitive to water deficits. The rate of leaf growth decreased linearly with predawn leaf water potential to reach zero at -1.5 MPa. Rate of leaf growth did not recover completely within the first three days after rewatering. Midday photosynthetic rates declined to 40% of those of well-watered seedlings at a predawn leaf water potential of -1.0 MPa and reached zero at -2.2 MPa. Photosynthetic rate recovered rapidly following rewatering and almost fully recovered by the second day after rewatering. All tissue water relations parameters, except the bulk modulus of elasticity, changed significantly as the soil dried and recovered completely by the third day after rewatering. Changes in osmotic pressure at full turgor of 0.4 MPa indicated considerable capacity for osmotic adjustment. However, because there was little osmotic adjustment until predawn leaf water potential fell below -1.5 MPa, this capacity would not have enhanced seedling growth, although it may have increased seedling survival. The sensitivity of photosynthesis and relative water content to water deficits suggests that greenhouse-grown E. marginata seedlings behave like mesophytic plants, even though E. marginata seedlings naturally grow in a drought-prone environment.     

Diurnal patterns of leaf photosynthesis, conductance and water potential at the top of a lowland rain forest canopy in Cameroon: measurements from the Radeau des Cimes

Diurnal patterns of leaf conductance, net photosynthesis and water potential of five tree species were measured at the top of the canopy in a tropical lowland rain forest in southwestern Cameroon. Access to the 40 m canopy was by a large canopy-supported raft, the Radeau des Cimes. The measurements were made under ambient conditions, but the raft altered the local energy balance at times, resulting in elevated leaf temperatures. Leaf water potential was equal to or greater than the gravitational potential at 40 m in the early morning, falling to values as low as -3.0 MPa near midday. Net photosynthesis and conductance were typically highest during midmorning, with values of about 10-12 micro mol CO(2) m(-2) s(-1) and 0.2-0.3 mol H(2)O m(-2) s(-1), respectively. Leaf conductance and net photosynthesis commonly declined through midday with occasional recovery late in the day. Photosynthesis was negatively related to leaf temperature above midday air temperature maxima. These patterns were similar to those observed in other seasonally droughted evergreen communities, such as Mediterranean-climate shrubs, and indicate that environmental factors may cause stomatal closure and limit photosynthesis in tropical rain forests during the midday period.     

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.     

Control of growth of juvenile leaves of Eucalyptus globulus: effects of leaf age

Biophysical variables influencing the expansion of plant cells (yield threshold, cell wall extensibility and turgor) were measured in individual Eucalyptus globulus leaves from the time of emergence until cessation of growth. Leaf water relations variables and growth rates were determined as relative humidity was changed on an hourly basis. Yield threshold and cell wall extensibility were estimated from plots of leaf growth rate versus turgor. Cell wall extensibility was also measured by the Instron technique, and yield threshold was determined experimentally both by stress relaxation in a psychrometer chamber and by incubation in a range of polyethylene glycol solutions. Once emerging leaves reached approximately 5 cm(2) in size, increases in leaf area were rapid throughout the expansive phase and varied little between light and dark periods. Both leaf growth rate and turgor were sensitive to changes in humidity, and in the longer term, both yield threshold and cell wall extensibility changed as the leaf aged. Rapidly expanding leaves had a very low yield threshold and high cell wall extensibility, whereas mature leaves had low cell wall extensibility. Yield threshold increased with leaf age.     

An analysis of elastic and plastic fruit growth of mango in response to various assimilate supplies

Changes in elastic and plastic components of mango (Mangifera indica L. cv 'Cogshall') fruit growth were analyzed with a model of fruit growth over time and in response to various assimilate supplies. The model is based on water relations (water potential and osmotic and turgor pressures) at the fruit level. Variation in elastic fruit growth was modeled as a function of the elastic modulus and variation in turgor pressure. Variation in plastic fruit growth was modeled using the Lockhart (1965) equation. In this model, plastic growth parameters (yield threshold pressure and cell wall extensibility) varied during fruit growth. Outputs of the model were diurnal and seasonal fruit growth, and fruit turgor pressure. These variables were simulated with good accuracy by the model, particularly the observed increase in fruit size with increasing availability of assimilate supply. Shrinkage was sensitive to the surface conductance of fruit peel, the elasticity modulus and the hydraulic conductivity of fruit, whereas fruit growth rate was highly sensitive to parameters linked to changes in wall extensibility and yield threshold pressure, regardless of the assimilate supply. According to the model, plastic growth was generally zero during the day and shrinkage and swelling were linked to the elastic behavior of the fruit. During the night, plastic and elastic growths were positive, resulting in fruit expansion.     

Adjustments in leaf water relations of mangrove (Avicennia germinans) seedlings grown in a salinity gradient

We used pressure-volume analysis and dewpoint hygrometry to determine leaf water relation parameters of mangrove (Avicennia germinans L.) seedlings grown at salinities of 0, 8, 20 and 32 per thousand. Seedlings responded to an increase in salinity from 0 to 32 per thousand by an increase in leaf succulence as reflected in an increase in leaf water content per unit area from 300 to 360 g m(-2). Additionally, osmotic potential at full turgor decreased from -2.3 to -3.5 MPa and osmotic potential at zero turgor decreased from -2.7 to -4.3 MPa. Cell elasticity decreased as salinity increased from 0 to 32 per thousand, as indicated by a progressive increase in volumetric modulus of elasticity from 19 to 27 MPa. Increased leaf succulence enabled leaves to sequester large amounts of solutes without adversely increasing cell osmotic pressure. On the other hand, osmotic adjustment facilitated turgor maintenance as water potential diminished. Salinity-induced decreases in tissue elasticity generated greater water potential differences between leaves and soil under saline conditions than under non-saline conditions.     

Osmotic potential of several hardwood species as affected by manipulation of throughfall precipitation in an upland oak forest during a dry year

Components of dehydration tolerance, including osmotic potential at full turgor (Psi(pio)) and osmotic adjustment (lowering of Psi(pio)), of several deciduous species were investigated in a mature, upland oak forest in eastern Tennessee. Beginning July 1993, the trees were subjected to one of three throughfall precipitation treatments: ambient, ambient minus 33% (dry treatment), and ambient plus 33% (wet treatment). During the dry 1995 growing season, leaf water potentials of all species declined to between -2.5 and -3.1 MPa in the dry treatment. There was considerable variation in Psi(pio) among species (-1.0 to -2.0 MPa). Based on Psi(pio) values, American beech (Fagus grandifolia Ehrh.), dogwood (Cornus florida L.), and sugar maple (Acer saccharum Marsh.) were least dehydration tolerant, red maple (A. rubrum L.) was intermediate in tolerance, and white oak (Quercus alba L.) and chestnut oak (Quercus prinus L.) were most tolerant. During severe drought, overstory chestnut oak and understory dogwood, red maple and chestnut oak displayed osmotic adjustment (-0.12 to -0.20 MPa) in the dry treatment relative to the wet treatment. (No osmotic adjustment was evident in understory red maple and chestnut oak during the previous wet year.) Osmotic potential at full turgor was generally correlated with leaf water potential, with both declining over the growing season, especially in species that displayed osmotic adjustment. However, osmotic adjustment was not restricted to species considered dehydration tolerant; for example, dogwood typically maintained high Psi(pio) and displayed osmotic adjustment to drought, but had the highest mortality rates of the species studied. Understory saplings tended to have higher Psi(pio) than overstory trees when water availability was high, but Psi(pio) of understory trees declined to values observed for overstory trees during severe drought. We conclude that Psi(pio) varies among deciduous hardwood species and is dependent on canopy position and soil water potential in the rooting zone.     

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.     

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.     

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.     

Water relations of loblolly pine seedlings from diverse geographic origins

Leaf conductance at three absolute humidity deficits (AHDs) (7, 14 and 21 g m(-3)), hydraulic conductance and components of tissue water potential were measured in one-year-old loblolly pine seedlings from six origins representing the geographic range of the species. Measurements were made on seedlings grown (a) with ample water (moist regime) and (b) with recurring severe drought (dry regime). However, all seedlings were well-watered prior to and during measurements. Seedlings grown in the moist regime had greater mean leaf conductances (0.30 versus 0.13 cm s(-1)) and greater responses to AHD than seedlings grown in the dry regime. They also exhibited greater hydraulic conductances (0.53 versus 0.35 microg cm(-2) s(-1) MPa(-1), less negative osmotic potentials (-1.45 versus -1.57 MPa) and higher relative water contents at turgor loss (0.72 versus 0.65). Seed source differences in water relations characteristics were detected only in seedlings grown in the moist regime. In these, trees from the three interior origins had greater mean leaf conductances than those from the three coastal sources (0.32 versus 0.28 cm s(-1)), but no differences in response to changing AHD were observed. Seedlings from North Carolina had lower osmotic potentials at turgor loss than those from Florida, Georgia or Texas. These differences in water relations characteristics are not clearly related to the observed greater survival ability of trees from interior origins compared with those from coastal origins.     

Phenotypic variation and quantitative trait locus identification for osmotic potential in an interspecific hybrid inbred F2 poplar pedigree grown in contrasting environments

Elucidation of the mechanisms of dehydration tolerance in poplar (Populus sp.) trees will permit development of biochemical and molecular indicators to identify dehydration-tolerant genotypes during genetic selection. The objectives of this study were to characterize the degree of phenotypic variation in osmotic potential (a determinant of dehydration tolerance), determine the relationship between osmotic potential at full turgor and relative growth rate, and identify quantitative trait loci (QTL) for osmotic potential in an advanced-generation, interspecific poplar pedigree established in contrasting environments. A three-generation, sib-mated black cottonwood (Populus trichocarpa Torr. & Gray) and eastern cottonwood (P. deltoides Bartr.) segregating F(2) family (Family 331) was analyzed at a dry site east of the Cascade Mountain Range (Boardman, OR) and at a wet site west of the mountains (Clatskanie, OR). At the Boardman site, 2-year-old trees (59 clones) were either irrigated everyday (wet) or every other day (dry), whereas 3- and 4-year-old trees (58 clones) at the Clatskanie site were unirrigated. At the Boardman site, the typically narrow range of osmotic potentials exhibited by grandparents and parents was greatly expanded in the F(2) population, spanning from -1.38 to -2.35 MPa under wet conditions, with a similar range under dry conditions (-1.40 to -2.15 MPa). Clones that had osmotic potentials < or = -1.90 MPa generally displayed full maintenance of stem relative growth rates under dry conditions in contrast to clones with osmotic potentials that were < or = -1.60 MPa, in which stem relative growth rates were reduced by an average of 38% in the dry treatment relative to the wet treatment. Although osmotic adjustments of 0.13 to 0.36 MPa were observed in nine out of 59 clones, adjustment typically occurred from relatively high baseline osmotic potentials. The range in osmotic potential at the wetter Clatskanie site at age three was higher (-1.27 to -1.84 MPa) and was further expanded the following year (-1.14 to -1.94 MPa), which had a wetter spring than the previous year, followed by a typically dry July. Seven QTL for osmotic potential were identified that each explained > 7.5% of the variation in osmotic potential. Given that four clones (7%) had osmotic potentials of -2.00 MPa or less and that QTL for osmotic potential have been identified, we suggest that there are opportunities to extend the limit of dehydration tolerance in Populus.     

Relationship of water status to vegetative growth and leaf gas exchange of peach (Prunus persica) trees on different rootstocks

We investigated relationships between tree water status, vegetative growth and leaf gas exchange of peach trees growing on different rootstocks under field conditions. Tree water status was manipulated by partially covering (0, approximately 30 and approximately 60%) the tree canopies on individual days and then evaluating the effects of tree water status on vegetative growth and leaf gas exchange. Early morning stem water potentials were approximately -0.4 MPa for trees in all treatments, but mean midday values ranged from -1.1 to -1.7 MPa depending on rootstock and canopy coverage treatment. Relative shoot extension growth rate, leaf conductance, transpiration rate and net CO2 exchange rate differed significantly among trees in the different rootstocks and canopy coverage treatments. Shoot extension growth rate, leaf conductance, leaf transpiration rate and leaf net CO2 exchange rate were linearly correlated with midday stem water potential. These relationships were independent of the rootstock and canopy coverage treatments, indicating that tree water relations are probably directly involved in the mechanism that imparts vegetative growth control by selected peach rootstocks.     

Clonal and seasonal differences in leaf osmotic potential and organic solutes of five hybrid poplar clones grown under field conditions

Leaf osmotic potential at full turgor (Psi(pio)) and the major solutes that contribute to osmotic potential were characterized in five hybrid poplar clones of Populus trichocarpa Torr. & Gray x P. deltoides Bartr. (TD) and P. deltoides x P. nigra L. (DN), growing under field conditions at two sites in eastern Washington and Oregon, USA. Trees were drip irrigated with 46, 76 or 137 cm of supplemental irrigation during each growing season. Trees at Wallula, WA, which were in their third growing season in 1994, were sampled twice a year for two years (1994 and 1995), and trees at Boardman, OR, which were in their second growing season in 1994, were sampled once a year for three years (1994-1996). At Wallula, the TD and DN clones exhibited lower predawn leaf water potentials in the 46-cm treatment than in the 137-cm treatment (-1.2 versus -0.7 MPa) during a hot, dry period in July 1994. Clone TD had a lower Psi(pio) than Clone DN (-1.67 versus -1.56 MPa) during the same period and the difference was also evident in 1995 (-1.81 versus -1.72 MPa) when trees were in their fourth growing season. There was also a significant treatment effect on Psi(pio) in Clone TD, with trees in the 46-cm treatment having lower Psi(pio) than trees in the 137-cm treatment in July 1994. At Boardman, Psi(pio) was generally high with no treatment differences during the 1994-96 samplings. The TD clones had significantly lower Psi(pio) than the DN clones in 1994 (-1.44 versus -1.36 MPa) and 1996 (-1.72 versus -1.54 MPa), but there was no difference between clones in 1995 (-1.40 versus -1.43 MPa). In 1995, at Wallula, osmotic adjustment in Clone TD was largely accounted for by an increase in sucrose, which constituted 70% of total organic solutes. Although the total concentration of free primary amino acids in this clone was 28% higher in trees in the 46-cm treatment than in trees in the 137-cm treatment, amino acids constituted only a small fraction of the total solute pool. Sixty-two percent of total solutes were inorganic ions in Clone TD compared to 52% in Clone DN, and potassium was the main ion constituting about 30% of total solutes and 50% of total ions. However, the clonal difference in Psi(pio) was not fully accounted for by the difference in solute concentration. Osmotic potential at full turgor declined over the growing season and with age. We conclude that, because the extent of osmotic adjustment exhibited by these clones was small, other drought resistance mechanisms contributed to the clonal differences in field performance.     

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.     

Gas exchange and water relations of Fraxinus americana affected by flurprimidol

Effects of flurprimidol on plant water relations and leaf gas exchange were investigated in one-year-old white ash (Fraxinus americana L.) seedlings subjected to soil water deficits. Flurprimidol (20 mg kg(-1) of soil equivalent) was applied to the soil surface of pot-grown seedlings after shoot growth was completed. Two months after flurprimidol application, water was withheld from one-half of the seedlings. Leaf water relations and gas exchange parameters were measured 5, 7, 10, 14, 18 and 22 days after withholding water. Under both irrigated and nonirrigated conditions, flurprimidol treatment resulted in reduced net CO(2) assimilation rate and transpirational water loss of seedlings as a result of decreased stomatal conductance. Consequently, flurprimidol-treated seedlings had higher leaf water potential and relative water content than untreated seedlings. Nonirrigated flurprimidol-treated seedlings also had greater turgor and sap osmolality and lower osmotic potential at full turgor than seedlings in the other treatments, indicating that flurprimidol increased osmotic adjustment. Under water-stress conditions, water use efficiency was lower and gas exchange efficiency was higher in flurprimidol-treated seedlings than in untreated seedlings, suggesting that flurprimidol treatment enhances survival of plants subjected to soil water deficits.     

Variation in water relations characteristics of terminal shoots of Port-Orford-cedar (Chamaecyparis lawsoniana) seedlings

We measured water relations attributes of the terminal shoots of 3-year-old Port-Orford-cedar (Chamaecyparis lawsoniana (A. Murr.) Parl.) seedlings that represented its geographic range. Pressure-volume curves were developed and osmotic potentials at full (psi(sf)) and zero turgor (psi(sz)), relative water content at zero turgor, and an index of tissue elasticity (IE) were calculated for 38 families during early, mid- and late summer at an inland nursery, and for 12 of these families during mid- and late summer at a coastal nursery. Compared with other conifer species, psi(sz) was high (-1.4 to -1.5 MPa) and declined in seedlings at both nurseries as the season progressed. Both IE and osmotic amplitude (psi(sf)-psi(sz)) increased during the season. Osmotic potential at zero turgor was lower and osmotic amplitude greater in seedlings at the inland nursery than at the coastal nursery. Correlations of water relations attributes with geographic location of the seed sources were weak and usually not significant. High elevation southern sources exhibited smaller differences in psi(sz) between nurseries than low elevation northern sources. The small differences in water relations attributes among sources and between nurseries suggest that some may be of marginal physiological importance; however, sources that produced larger seedlings appeared to be less desiccation tolerant. We conclude that, when moving genotypes during reforestation, decisions based on patterns in tree size and timing of growth will account for these small differences in water relations.     

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.