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.     

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.     

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.     

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.     

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.     

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).     

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.     

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.     

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).     

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.     

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.     

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.     

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.     

Hydraulic and stomatal adjustment of Norway spruce trees to environmental stress

A study of how the water conducting systems of 30-50-year-old Norway spruce (Picea abies (L.) Karst.) trees growing at three sites adjust to shade and waterlogging indicated that water relations characteristics varied with the life histories of the trees. Xylem was more efficient at conducting water and stomata were more sensitive to atmospheric evaporative demand in trees subjected to favorable growth conditions (control trees) than in trees growing in shade or waterlogged conditions. At the same soil water availability, shade-grown trees suffered more severely from water deficit than control trees. Under conditions of high atmospheric vapor pressure deficit, foliage of shade-grown trees exhibited low water potentials, as a result of low hydraulic conductance of the vascular system and inefficient stomatal control. Because of the increased internal resistance to water flow, more negative leaf water potentials (Psi(x)) must be reached to provide an adequate water supply to the foliage. It is concluded that dynamic water stress is one of the main causes of the continuing growth retardation in suppressed Norway spruce trees after their release from the overstory. Trees growing in waterlogged soil (bog-grown trees) were characterized by weak stomatal control, resulting in large water losses from the foliage. Although bog-grown trees exhibited uneconomical water use, they possessed mechanisms (e.g., osmotic adjustment) that allowed leaves to tolerate low Psi(x) while stomata remained open. Under conditions of sufficient soil water availability and moderate atmospheric vapor pressure deficit, soil-to-leaf conductance was highest in bog-grown trees (1.45 +/- 0.06 mmol m(-2) s(-1) MPa(-1)), followed by control and shade-grown trees (1.04 +/- 0.04 and 0.77 +/- 0.05 mmol m(-2) s(-1) MPa(-1), respectively). The lowest soil-to-leaf conductance (0.45 +/- 0.04 mmol m(-2) s(-1) MPa(-1)) was recorded in control trees at high atmospheric evaporative demand, and was probably caused by tracheid cavitation.     

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.     

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

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

Foliar Responses of Understorey Abies lasiocarpa to Different Degrees of Release Cutting in a Betula papyrifera and Conifer Mixed Species Stand

Foliar responses of subalpine fir [Abies lasiocarpa (Hook.) Nutt.] to thinning were studied in a 35-yr-old mixed stand of paper birch (Betula papyrifera Marsh.) and conifers. The stand regenerated naturally after a wildfire with a canopy dominated by paper birch (average height 9.8 m) and an understorey dominated by subalpine fir (average height 1.6 m). The stand was thinned to four densities of birch: 0, 600 and 1200 stems ha-¹ and control (unthinned at 2300-6400 stems ha-¹) in the autumn of 1995. The understorey conifers, mainly subalpine fir, were thinned to 1200 stems ha-¹. The study used a completely randomized split-plot design. Three sample trees were systematically selected from each treatment replicate and each tree stratum (upper, intermediate and lower understorey). One-year-old and older age class needles were collected from one south-facing branch within the fifth whorl from the tree top. Thinning of paper birch significantly (p <0.001) increased leaf area and dry weight per 100 needles for intermediate and short trees except in the 0 birch treatment. Understorey subalpine fir trees in 600 stems ha¹ birch (T3) had the largest leaf area and leaf dry weight per 100 1-yr-old needles. Specific leaf area (SLA) decreased from unthinned (T1) to 0 birch (T4). Lower understorey trees had the largest SLA. One-year-old needles had significantly higher N, P and K concentrations in all the thinning treatments. These responses are consistent with the shade tolerance of subalpine fir. The results suggest that when managing a paper birch-conifers mixed-wood forest it may be of benefit to understorey conifers to leave a birch canopy as a nursing crop.     

Water Relations in Sierra Nevada Mixed Conifer: Wildfire Effects and Species Distinctions

An investigation of wildfire effects on water relations of Jeffrey pine (Pinus jeffreyi Grev. & Balf.) along with species comparisons of the water relations of unburned specimens of this pine to those of unburned white fir (Abies concolor var. lowiana [Gord.] Lemm.) were conducted in a mixed conifer stand located in the Lake Tahoe Basin. Xylem water potentials were significantly lower in burned than in the unburned Jeffrey pine as measured in predawn, midday, and evening periods distributed over more than two postfire growing seasons, while soil water potentials were lower in burned than in unburned stand portions during the drier parts of the growing season, but the reverse proved true during the wetter part. Diurnal fluctuation in bole diameter, a measure of stored water recharge capacity, was largely unaffected by wildfire, however. Xylem water potentials were consistently lower in unburned white fir than in unburned Jeffrey pine and DBH fluctuation was often lower in the fir than in the pine as well, but soil water potentials associated with unburned subject trees did not differ significantly between the two species. Demonstrated here are ecophysiological alterations that occur in Sierra Nevada mixed conifer due to wildfire and shifting species composition.     

Impacts of hemlock looper defoliation on growth and survival of balsam fir, black spruce and white birch in Newfoundland, Canada

Hemlock looper (Lambdina fiscellaria fiscellaria (Guen.)) is an economically damaging defoliator that undergoes periodic outbreaks in Newfoundland, Canada. It defoliates and causes extensive tree mortality to its primary host, balsam fir (Abies balsamea [L.] Mill.). We quantified tree survival using data from permanent sample plots (PSPs) and growth reduction or release using dendrochronology, and related these impacts to defoliation severity determined from annual aerial defoliation survey data. Such impact relationships are necessary as a fundamental input to a Decision Support System. Growth and survival of balsam fir, black spruce (Picea mariana (Mill.) B.S.P.) and white birch (Betula papyrifera Marsh.) were assessed from 1996 to 2008 in 48 Newfoundland Forest Service PSPs, selected based on four classes of defoliation severity. Two years of severe (71-100%) defoliation resulted in almost complete mortality for balsam fir, 10 years after defoliation, whereas survival was 70-80% for black spruce and white birch. Lower defoliation severity (1-2 years of moderate (31-70%) or 1 year of severe) resulted in approximately 60% survival for balsam fir and no reduction in survival for black spruce and white birch. Maximum growth reduction of balsam fir was 10-15% with 1 year of moderate-severe defoliation, 35-40% with 2 years of moderate defoliation, and about 50% with 2 years of severe defoliation. Growth recovered to pre-defoliation rates 5 years after defoliation ceased in all severity classes. Growth reduction and recovery of black spruce were more variable and lower than for balsam fir, and white birch exhibited only minor (<10%) growth reduction during the defoliation year or 1 year after defoliation. Control measures should focus on avoiding severe defoliation for two consecutive years.     

Influence of soil texture on hydraulic properties and water relations of a dominant warm-desert phreatophyte

We investigated hydraulic constraints on water uptake by velvet mesquite (Prosopis velutina Woot.) at a site with sandy-loam soil and at a site with loamy-clay soil in southeastern Arizona, USA. We predicted that trees on sandy-loam soil have less negative xylem and soil water potentials during drought and a lower resistance to xylem cavitation, and reach E(crit) (the maximum steady-state transpiration rate without hydraulic failure) at higher soil water potentials than trees on loamy-clay soil. However, minimum predawn leaf xylem water potentials measured during the height of summer drought were significantly lower at the sandy-loam site (-3.5 +/- 0.1 MPa; all errors are 95% confidence limits) than at the loamy-clay site (-2.9 +/- 0.1 MPa). Minimum midday xylem water potentials also were lower at the sandy-loam site (-4.5 +/- 0.1 MPa) than at the loamy-clay site (-4.0 +/- 0.1 MPa). Despite the differences in leaf water potentials, there were no significant differences in either root or stem xylem embolism, mean cavitation pressure or Psi(95) (xylem water potential causing 95% cavitation) between trees at the two sites. A soil-plant hydraulic model parameterized with the field data predicted that E(crit) approaches zero at a substantially higher bulk soil water potential (Psi(s)) on sandy-loam soil than on loamy-clay soil, because of limiting rhizosphere conductance. The model predicted that transpiration at the sandy-loam site is limited by E(crit) and is tightly coupled to Psi(s) over much of the growing season, suggesting that seasonal transpiration fluxes at the sandy-loam site are strongly linked to intra-annual precipitation pulses. Conversely, the model predicted that trees on loamy-clay soil operate below E(crit) throughout the growing season, suggesting that fluxes on fine-textured soils are closely coupled to inter-annual changes in precipitation. Information on the combined importance of xylem and rhizosphere constraints to leaf water supply across soil texture gradients provides insight into processes controlling plant water balance and larger scale hydrologic processes.