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.     

Solute accumulation of chestnut oak and dogwood leaves in response to throughfall manipulation of an upland oak forest

To determine the biochemical basis of osmotic adjustment, seasonal and treatment differences in foliar water- soluble organic solutes and inorganic ions were investigated for two hardwood species that exhibited osmotic adjustment in a Throughfall Displacement Experiment at the Walker Branch Watershed near Oak Ridge, Tennessee. Leaf samples of overstory and understory chestnut oak (Quercus prinus L.) and understory dogwood (Cornus florida L.) were collected during the 1994 (wet) and 1995 (dry) growing seasons from each of three treatments: dry (-33% throughfall), ambient (control) and wet (+33% throughfall). Foliar soluble carbohydrates and organic acids were measured by gas chromatography-mass spectrometry. During May 1994, when the demand for sucrose was greatest, dogwood accumulated small amounts of glucose, quinic acid and Mg2+, offsetting a decline in nitrate concentration. As the mild drought continued and tree growth slowed, there was a significant accumulation of sucrose in dogwood in the dry treatment in June, and a trend toward increased K+. In overstory chestnut oak in the dry treatment over the same period, there were significant accumulations of fructose, glucose and K+, and a trend toward increased quinic acid accumulation. Sucrose did not become a key osmotically active compound in chestnut oak until August 1994. In 1995, with the exception of understory chestnut oak, there was no accumulation of K+ in either species. During the severe drought of 1995, monosaccharides, particularly glucose and fructose, accounted for most of the osmotic adjustment in both species. Among solutes, glucose constituted the largest accumulation in dogwood in the dry treatment in August 1995, followed by fructose and sucrose. There was only a moderate increase in solutes in chestnut oak trees in 1995, with fructose and glucose constituting over 50% of the predicted solute accumulation in July. Both species accumulated a wider array of solutes during the dry year than during the wet year, but treatment differences in solute accumulation in chestnut oak were partially limited by drought. The greater dehydration tolerance of chestnut oak than dogwood was evident in the higher baseline concentrations of organic solutes and the greater array of solutes accumulated in response to drought.     

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.     

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.     

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.     

Water relations and gas exchange of Acer saccharum seedlings in contrasting natural light and water regimes

Field measurements were made of leaf photosynthesis (A), stomatal conductance (g) and leaf water relations for sugar maple (Acer saccharum Marsh.) seedlings growing in a forest understory, small gap or large clearing habitat in southwestern Wisconsin, USA. Predawn water status, leaf gas exchange and plasticity in field and laboratory water relations characteristics were compared among contrasting light environments in a wet year (1987) and a dry year (1988) to evaluate possible interactions between light and water availability in these habitats. Leaf water potentials (Psi(leaf)) at predawn and midday were lower for clearing than gap or understory seedlings. Acclimation of tissue osmotic potentials to light environment was observed among habitats but did not occur within any of the habitats in response to prolonged drought. During a summer drought in 1988, decreases in daily maximum g (g(max)) and maximum A (A(max)) in clearing seedlings were correlated with predawn Psi(leaf), which reached a seasonal minimum of -2.0 MPa. Under well-watered conditions, diurnal fluctuations in Psi(leaf) of up to 2.0 MPa in clearing seedlings occurred along with large midday depressions of A and g. In a wet year, strong stomatal responses to leaf-to-air vapor pressure difference (VPD) in sunny habitats were observed over nine diurnal courses of gas exchange measurements on seedlings in a gap and a clearing. Increasing stomatal limitations to photosynthesis appeared to be responsible for the reduction in A at high VPD for clearing seedlings. In understory seedlings, however, low water-use efficiency and development of leaf water deficits in sunflecks was related to reduced stomatal limitations to photosynthesis relative to seedlings in sunny habitats. Predawn Psi(leaf) and VPD appear to be important factors limiting carbon assimilation in sugar maple seedlings in light-saturating irradiances, primarily through stomatal closure. The overall results are consistent with the idea that sugar maple seedlings exhibit "conservative" water use patterns and have low drought tolerance. Leaf water relations and patterns of water use should be considered in studies of acclimation and species photosynthetic performance in contrasting light environments.     

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.     

Relationships between advance oak regeneration and biotic and abiotic factors

Relationships between advance regeneration of four tree species (red maple (Acer rubrum L.), white oak (Quercus alba L.), chestnut oak (Q. montana Willd.) and northern red oak (Q. rubra L.)) and biotic (non-tree vegetation and canopy composition) and abiotic (soil series and topographic variables) factors were investigated in 52, mature mixed-oak stands in the central Appalachians. Aggregate height was used as a composite measure of regeneration abundance. Analyses were carried out separately for two physiographic provinces. Associations with tree regeneration were found for all biotic and abiotic factors both in partial models and full models. Red maple was abundant on most of the sites, but high red maple abundance was commonly associated with wet north-facing slopes with little or no cover of mountain-laurel (Kalmia latifolia L.) and hay-scented fern (Dennstaedtia punctilobula (Michx.) Moore). Regeneration of the three oak species was greatly favored by the abundance of overstory trees of their own kind. White oak regeneration was most abundant on south-facing, gentle, lower slopes with soils in the Buchanan series. Chestnut oak regeneration was more common on south-facing, steep upper slopes with stony soils. There was a positive association between chestnut oak and huckleberry (Gaylussacia baccata (Wangh.) Koch) cover classes. Northern red oak was more abundant on north-facing wet sites with Hazleton soil, and was associated with low occurrence of mountain-laurel and hay-scented fern.     

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.     

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.     

Forest regeneration composition and development in upland, mixed-oak forests

Advance regeneration in 52 mature mixed-oak stands was analyzed and described. Red maple (Acer rubrum L.) was the most abundant species in the study area. Among oak (Quercus) species, northern red oak (Q. rubra L.) was the most abundant within the Allegheny Plateau physiographic province, whereas chestnut oak (Q. montana L.) was the most abundant within the Ridge and Valley physiographic province. Sixteen stands, for which data are available through the fourth growing season following harvest, were used to describe stand development. Cumulative height, a composite measure of size and density, was used to describe early stand development. Black gum (Nyssa sylvatica Marsh.) and black birch (Betula lenta L.) had dramatic increases in stand density and cumulative height after overstory removal. Cumulative height of northern red oak and chestnut oak showed a faster positive response to overstory removal than red maple. Oak retained its dominance in cumulative height for at least 4 years after harvest. Red maple nevertheless remained the most abundant tree species after overstory removal. Our results suggest that the principal advantage of red maple regeneration is its ability to accumulate in large numbers prior to harvest.     

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.     

Shoot water relations of mature black spruce families displaying a genotype x environment interaction in growth rate. I. Family and site effects over three growing seasons

Pressure-volume curves were determined for black spruce (Picea mariana (Mill.) BSP) trees from four full-sib families. During the first two years, trees were measured from a plantation on a dry site. In the third year, trees were sampled from the dry site and a wet site. Diurnal measurements of shoot water potential allowed in situ shoot turgor to be estimated in addition to standard water relations traits. Over all years, Female 59 progeny displayed lower osmotic potentials at saturation (Psi(sat)) than Female 63 progeny. Genetic differences in Psi(sat) were similar on both the dry and wet sites. Modulus of elasticity (epsilon) was greater for Female 59 progeny than for Female 63 progeny, producing a compensatory effect resulting in no genetic or site differences in osmotic potential at turgor loss point (Psi(tlp)) or relative water content at turgor loss point (RWC(tlp)). Mean and predawn shoot turgor pressures (P(x) and P(pd)) were higher for Female 59 progeny than for Female 63 progeny and higher at the wet site than the dry site. Genotype x environment trends were observed; compared to Female 63 progeny, Female 59 progeny displayed 9.8 and 5.1% higher P(pd) on the dry and wet sites, respectively, and 3.4 and 9.8% greater P(pd) values in wet and dry years, respectively. Tree volume growth showed no relationship to Psi(tlp) or RWC(tlp), but was correlated with Psi(sat) and P(x); however, the strongest correlation was with P(pd) (r = 0.90).     

A comparison of root growth dynamics of silver maple and flowering dogwood in compacted soil at differing soil water contents

Many bottomland tree species are tolerant of compacted soil and perform well in urban environments; however, the mechanism underlying this tolerance is unknown. Increased soil water content has been shown to alleviate some of the effects of soil compaction on plant growth, presumably because increasing soil water reduces soil strength. We hypothesized that tree species tolerant of very wet soils would have opportunities for root growth in compacted soil when high soil water contents reduced soil strength, whereas species intolerant of bottomland conditions would not. We tested this hypothesis on flowering dogwood (Cornus florida L.), a mesic species intolerant of inundation, and silver maple (Acer saccharinum L.), a bottomland species. Seedlings of both species were grown in pots for 21 and 30 days, respectively, in a growth chamber in native loam soil maintained at various combinations of soil strength and soil water tension. Downward root growth rate decreased in response to increasing soil strength in both species. At low soil strength (0.6 MPa), downward root growth rate of dogwood seedlings slowed when soil was either excessively wet or dry, whereas root growth rate of silver maple seedlings increased linearly with soil water content. In moderately compacted soil (1.5 g cm(-3) bulk density), silver maple seedlings had greater root growth rate, root length per plant, and ratio of root length to root dry weight in wet soil (0.006 MPa soil water tension) than in moist and dry soils (0.026 and 0.06 MPa, respectively), even though mean oxygen diffusion rate (ODR) was only 0.28 &mgr;g cm(-2) (SE = 0.05). No such effect was detected in highly compacted soil (1.7 g cm(-3) bulk density) in either species. Mean ODR showed a weak positive correlation with soil water tension (r = 0.40, P = 0.07), but was unrelated to soil strength. We conclude that silver maple roots can grow in moderately compacted soil when high soil water content decreases soil strength, whereas dogwood is unable to take advantage of this opportunity.     

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.     

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.     

Seasonal patterns of leaf gas exchange and water relations in dry rain forest trees of contrasting leaf phenology

Diurnal and seasonal patterns of leaf gas exchange and water relations were examined in tree species of contrasting leaf phenology growing in a seasonally dry tropical rain forest in north-eastern Australia. Two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., and two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret. were studied. The deciduous species had higher specific leaf areas and maximum photosynthetic rates per leaf dry mass in the wet season than the evergreens. During the transition from wet season to dry season, total canopy area was reduced by 70-90% in the deciduous species and stomatal conductance (g(s)) and assimilation rate (A) were markedly lower in the remaining leaves. Deciduous species maintained daytime leaf water potentials (Psi(L)) at close to or above wet season values by a combination of stomatal regulation and reduction in leaf area. Thus, the timing of leaf drop in deciduous species was not associated with large negative values of daytime Psi(L) (greater than -1.6 MPa) or predawn Psi(L) (greater than -1.0 MPa). The deciduous species appeared sensitive to small perturbations in soil and leaf water status that signalled the onset of drought. The evergreen species were less sensitive to the onset of drought and g(s) values were not significantly lower during the transitional period. In the dry season, the evergreen species maintained their canopies despite increasing water-stress; however, unlike Eucalyptus species from northern Australian savannas, A and g(s) were significantly lower than wet season values.     

Effect of gap size on mid-rotation stand structure and species composition in a naturally regenerated mixed broadleaf forest

This paper is an assessment of the effect of gap size on stand structure and species composition 48 years following treatment in a mixed broadleaf upland forest. Established in 1960, the study tests three circular openings, 15.2 m (0.02 ha), 45.7 m (0.16 ha), and 76.2 m (0.46 ha). Forty-eight years following treatment (2008) basal area, top height, and quadratic mean diameter were significantly lower in 15.2 m openings. Maple (Acer spp.) species had the highest mean importance value across treatments (0.40). Trends suggest that species composition of dominant and codominant trees among opening sizes may have been influenced by shade tolerance adaptations of the species groups present. Whereas 15.2 m openings were dominated by shade tolerant maple species, 45.7 and 76.2 m openings produced a mixture of commercial species including shade intolerant species such as yellow-poplar (Liriodendron tulipifera L.), trees of intermediate shade tolerance like oak (Quercus spp.), and shade tolerant maple. Data further suggest the density of overstory oak was highest in the intermediate opening size (45.7 m), while yellow-poplar increased in the larger opening sizes. Evaluation of species shifts between 1981 and 2008 showed that relative basal area of maple increased across all treatments. Relative basal areas from 45.7 to 76.2 m openings suggest declines in yellow-poplar and other non-commercial species were balanced by increases in oak and maple.     

Response of gas exchange to water stress in seedlings of woody angiosperms

Responses of net photosynthesis (A), leaf conductance to water vapor (g(wv)) and instantaneous water use efficiency (WUE) to decreasing leaf and soil water potentials (Psi(l), Psi(s)) were studied in three-month-old white oak (Quercus alba L.), post oak (Q. stellata Wangenh.), sugar maple (Acer saccharum Marsh.), and black walnut (Juglans nigra L.) seedlings. Quercus seedlings had the highest A and g(wv) when plants were well watered. As the soil was allowed to dry, both A and g(wv) decreased; however, trace amounts of A were observed at a Psi(l) as low as -2.9 MPa in Q. stellata and -2.6 MPa in Q. alba and A. saccharum. Photosynthesis was not measurable at Psi(l) lower than -2.2 MPa in J. nigra and water stress-induced leaflet senescence was observed in this species. Within each species, g(wv) showed a similar relationship to soil and leaf Psi, but the response to Psi(l) was shifted to more negative values by 1.2 to 1.6 MPa. As Psi(s) declined below -1 MPa, the difference between soil and leaf Psi diminished because of the suppression of transpiration. There was no indication that Psi(s) had a more direct influence on g(wv) than did Psi(l). Water use efficiency showed an initial increase as the soil dried, followed by a decline under severe water stress. Water use efficiency was highest in J. nigra, intermediate in Quercus species and lowest in A. saccharum. There was an evident relationship between gas exchange characteristics and natural distribution in these species, with the more xeric species showing higher A and g(wv) under both well-watered and water-stressed conditions. There was no trend toward increased efficiency of water use in the more xeric species.