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.     

Xylem cavitation and loss of hydraulic conductance in western hemlock following planting

Following planting, western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedlings experience water stress and declining xylem pressure potential (Psi(x)). Low Psi(x) can result in xylem cavitation and embolism formation, causing a decline in hydraulic conductance. This study focused on the relationship between Psi(x), xylem cavitation and transpiration (E) of newly planted seedlings. Leaf specific hydraulic conductance (k(AB)) declined from 0.56 to 0.09 mmol m(-2) s(-1) MPa(-1) over a 9-day period. Stomatal conductance (g(s)) declined from 143.5 to 39.15 mmol m(-2) s(-1) over the same period without an associated change in environmental conditions. A vulnerability profile indicated a 30% loss in hydraulic conductivity when seedlings experienced a Psi(x) between -2.5 and -3.0 MPa. A Psi(x) of -4.0 MPa led to a complete loss of conductivity. We conclude that following planting, western hemlock seedlings often experience Psi(x) values that are low enough to cause xylem cavitation and a decline in k(AB).     

Effects of ring-porous and diffuse-porous stem wood anatomy on the hydraulic parameters used in a water flow and storage model

Calibration of a recently developed water flow and storage model based on experimental data for a young diffuse-porous beech tree (Fagus sylvatica L.) and a young ring-porous oak tree (Quercus robur L.) revealed that differences in stem wood anatomy between species strongly affect the calibrated values of the hydraulic model parameters. The hydraulic capacitance (C) of the stem storage tissue was higher in oak than in beech (939.8 versus 212.3 mg MPa(-1)). Model simulation of the elastic modulus (epsilon) revealed that this difference was linked to the higher elasticity of the stem storage tissue of oak compared with beech. Furthermore, the hydraulic resistance (R (x)) of beech was about twice that of oak (0.1829 versus 0.1072 MPa s mg(-1)). To determine the physiological meaning of the R (x) parameter identified by model calibration, we analyzed the stem wood anatomy of the beech and oak trees. Calculation of stem specific hydraulic conductivity (k (s)) of beech and oak with the Hagen-Poiseuille equation confirmed the differences in R (x) predicted by the model. The contributions of different vessel diameter classes to the total hydraulic conductivity of the xylem were calculated. As expected, the few big vessels contributed much more to total conductivity than the many small vessels. Compared with beech, the larger vessels of oak resulted in a higher k (s) (10.66 versus 4.90 kg m(-1) s(-1) MPa(-1)). The calculated ratio of k (s) of oak to beech was 2, confirming the R (x) ratio obtained by model calibration. Thus, validation of the R (x) parameter of the model led to identification of its physiological meaning.     

Quantifying flooding effects on hardwood seedling survival and growth for bottomland restoration

Growing interest worldwide in bottomland hardwood restoration necessitates improved ecological understanding of flooding effects on forest tree seedlings using methodology that accurately reflects field conditions. We examined hardwood seedling survival and growth in an outdoor laboratory where the timing, depth, duration, and flow rate of flood water can be carefully controlled while simulating natural soil conditions occurring in floodplains. Flooding treatments were initiated in mid-May and included partial inundation (15–20?cm) during the growing season for 5-week flowing, 5-week standing, 3-week flowing, and control. We monitored the vigor, survival, and growth (changes in basal diameter and stem length) of six hardwood species representing a wide range in expected flood tolerance including eastern cottonwood (Populus deltoides Bartr. Ex Marsh.), pin oak (Quercus palustris Muenchh.), swamp white oak (Q. bicolor Willd.), bur oak (Q. macrocarpa Michx.), black walnut (Juglans nigra L.), and pecan [Carya illinoensis (Wangenh.) K. Koch]. All stock was 1-0 bareroot except that cuttings were used for eastern cottonwood. Five species—eastern cottonwood, bur oak, swamp white oak, pin oak, and pecan—exhibited high survival probabilities (>0.62 for cottonwood; >0.77 for the others) regardless of flood treatment. But of the survivors, only eastern cottonwood and swamp white oak maintained positive growth and healthy green foliage. Despite high survival, bur oak and pin oak suffered stem growth losses and exhibited chlorotic foliage in flood treatments suggesting greater vulnerability to other abiotic or biotic stresses if outplanted on flood-prone sites. Pecan also suffered stem dieback in controls suggesting vulnerability to competition and browsing when outplanted despite high survival after flooding. Our quantitative data helps to confirm and/or refine previously published qualitative flood tolerance ratings for these species, and describes operation of an in situ outdoor flood experiment laboratory that may prove effective in guiding future flood tolerance research.     

Vulnerability to drought-induced cavitation of riparian cottonwoods in Alberta: a possible factor in the decline of the ecosystem?

Vulnerability of xylem to loss of hydraulic conductivity caused by drought-induced cavitation was determined for three riparian cottonwood species in Lethbridge, Alberta: Populus deltoides Bartr., P. balsamifera L., and P. angustifolia James. These species suffered 50% loss of hydraulic conductivity in one-year-old stem segments when xylem pressure potential fell to -0.7 MPa for P. deltoides and -1.7 MPa for P. balsamifera and P. angustifolia, making them the three most vulnerable tree species reported so far in North America. The possible contribution of drought-induced xylem dysfunction to the decline of riparian ecosystems in dammed rivers is discussed.     

Water use and carbon exchange of red oak- and eastern hemlock-dominated forests in the northeastern USA: implications for ecosystem-level effects of hemlock woolly adelgid

Water use and carbon exchange of a red oak-dominated (Quercus rubra L.) forest and an eastern hemlock-dominated (Tsuga canadensis L.) forest, each located within the Harvard Forest in north-central Massachusetts, were measured for 2 years by the eddy flux method. Water use by the red oak forest reached 4 mm day(-1), compared to a maximum of 2 mm day(-1) by the eastern hemlock forest. Maximal carbon (C) uptake rate was also higher in the red oak forest than in the eastern hemlock forest (about 25 versus 15 micromol m(-2) s(-1)). Sap flux measurements indicated that transpiration of red oak, and also of black birch (Betula lenta L.), which frequently replaces eastern hemlock killed by hemlock woolly adelgid (Adelges tsugae Annand.), were almost twice that of eastern hemlock. Despite the difference between species in maximum summertime C assimilation rate, annual C storage of the eastern hemlock forest almost equaled that of the red oak forest because of net C uptake by eastern hemlock during unusually warm fall and spring weather, and a near-zero C balance during the winter. Thus, the effect on C storage of replacing eastern hemlock forest with a forest dominated by deciduous species is unclear. Carbon storage by eastern hemlock forests during fall, winter and spring is likely to increase in the event of climate warming, although this may be offset by C loss during hotter summers. Our results indicate that, although forest water use will decrease immediately following eastern hemlock mortality due to the hemlock woolly adelgid, the replacement of eastern hemlock by deciduous species such as red oak will likely increase summertime water use over current rates in areas where hemlock is a major forest species.     

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.     

Influence of overstory density on ecophysiology of red oak (Quercus rubra) and sugar maple (Acer saccharum) seedlings in central Ontario shelterwoods

A field experiment was established in a second-growth hardwood forest dominated by red oak (Quercus rubra L.) to examine the effects of shelterwood overstory density on leaf gas exchange and seedling water status of planted red oak, naturally regenerated red oak and sugar maple (Acer saccharum Marsh.) seedlings during the first growing season following harvest. Canopy cover of uncut control stands and moderate and light shelterwoods averaged 97, 80 and 49%, respectively. Understory light and vapor pressure deficit (VPD) strongly influenced gas exchange responses to overstory reduction. Increased irradiance beneath the shelterwoods significantly increased net photosynthesis (P(n)) and leaf conductance to water vapor (G(wv)) of red oak and maple seedlings; however, P(n) and G(wv) of planted and naturally regenerated red oak seedlings were two to three times higher than those of sugar maple seedlings in both partial harvest treatments, due in large part to decreased stomatal limitation of gas exchange in red oak as a result of increased VPD in the shelterwoods. In both species, seedling water status was higher in the partial harvest treatments, as reflected by the higher predawn leaf water potential and seedling water-use efficiency in seedlings in shelterwoods than in uncut stands. Within a treatment, planted and natural red oak seedlings exhibited similar leaf gas exchange rates and water status, indicating little adverse physiological effect of transplanting. We conclude that the use of shelterwoods favors photosynthetic potential of red oak over sugar maple, and should improve red oak regeneration in Ontario.     

Groundwater and Light Conditions as Factors in the Survival of Pedunculate Oak (Quercus Robur L.) Seedlings

A noticeable decline and physiological weakening of pedunculate oak (Quercus robur L.) has been recorded in the past decade in lowland parts of Slovenia. The triggers were dry climate, unfavourable precipitation patterns and human influence that brought about changes in watercourses and water tables. Regeneration and future management of this tree species is therefore questionable. To define a critical groundwater table which would permit successful regeneration, planted and naturally grown seedlings were compared in two forests differing in the degree of decline and physiological weakness of adult pedunculate oak trees. Comparison of stomatal conductance and pre-dawn water potential (PWP) showed differences of seedlings between and within forest complexes. Stomatal conductance dropped below 100 mmol m⁻² s⁻¹ at PWP values below −1.0 MPa, while closure of stomata was caused by a PWP between −1.6 and −1.9 MPa. Increased water stress equalized photosynthetic yield in both natural and planted oaks below −1.4 MPa.     

Oak seedling growth and ectomycorrhizal colonization are less in eastern hemlock stands infested with hemlock woolly adelgid than in adjacent oak stands

Invasive, non-indigenous, phytophagous insects have caused widespread declines in several dominant tree species. The decline in dominant tree species may lead to cascading effects on other tree and microbial species and their interactions, affecting forest recovery following the decline. In the eastern USA, eastern hemlock (Tsuga canadensis (L.) Carr) is declining because of infestation by the hemlock woolly adelgid (HWA; Adelges tsugae Annand). Northern red oak (Quercus rubra L.) is a common replacement species in declining hemlock stands, but reduced mycorrhizal inoculum potential in infested hemlock stands may cause oak to grow more slowly compared with oak in oak stands. We grew red oak seedlings for one growing season in declining hemlock-dominated stands infested with HWA and in adjacent oak-dominated stands. Ectomycorrhizal root tip density and morphotype richness in soil cores were 63 and 27% less, respectively, in declining hemlock stands than in oak stands. Similarly, ectomycorrhizal percent colonization and morphotype richness on oak seedlings were 33 and 30% less, respectively, in declining hemlock stands than in oak stands. In addition, oak seedlings in declining hemlock stands had 29% less dry mass than oak seedlings in oak stands. Analysis of covariance indicated that morphotype richness could account for differences in oak seedling dry mass between declining hemlock stands and oak stands. Additionally, oak seedling dry mass in declining hemlock stands significantly decreased with decreasing ectomycorrhizal percent colonization and morphotype richness. These results suggest that oak seedling growth in declining hemlock stands is affected by reduced ectomycorrhizal inoculum potential. Further, the rate of forest recovery following hemlock decline associated with HWA infestation may be slowed by indirect effects of HWA on the growth of replacement species, through effects on ectomycorrhizal colonization and morphotype richness.     

Cavitation in dehydrating xylem of Picea abies: energy properties of ultrasonic emissions reflect tracheid dimensions

Ultrasonic emission (UE) testing is used to analyse the vulnerability of xylem to embolism, but the number of UEs often does not sufficiently reflect effects on hydraulic conductivity. We monitored the absolute energy of UE signals in dehydrating xylem samples hypothesizing that (i) conduit diameter is correlated with UE energy and (ii) monitoring of UE energy may enhance the utility of this technique for analysis of xylem vulnerability. Split xylem samples were prepared from trunk wood of Picea abies, and four categories of samples, derived from mature (I: earlywood, II: 30-50% latewood, III: >50% latewood) or juvenile wood (IV: earlywood) were used. Ultrasonic emissions during dehydration were registered and anatomical parameters (tracheid lumen area, number per area) were analysed from cross-sections. Attenuation of UE energy was measured on a dehydrating wood beam by repeated lead breaks. Vulnerability to drought-induced embolism was analysed on dehydrating branches by hydraulic, UE number or UE energy measurements. In split samples, the cumulative number of UEs increased linearly with the number of tracheids per cross-section, and UE energy was positively correlated with the mean lumen area. Ultrasonic emission energies of earlywood samples (I and IV), which showed normally distributed tracheid lumen areas, increased during dehydration, whereas samples with latewood (II and III) exhibited a right-skewed distribution of lumina and UE energies. Ultrasonic emission energy was hardly influenced by moisture content until ～40% moisture loss, and decreased exponentially thereafter. Dehydrating branches showed a 50% loss of conductivity at -3.6 MPa in hydraulic measurements and at -3.9 and -3.5 MPa in UE analysis based on cumulative number or energy of signals, respectively. Ultrasonic emission energy emitted by cavitating conduits is determined by the xylem water potential and by the size of element. Energy patterns during dehydration are thus influenced by the vulnerability to cavitation, conduit size distribution as well as attenuation properties. Measurements of UE energy may be used as an alternative to the number of UEs in vulnerability analysis.     

Leaf gas exchange characteristics differ among Sonoran Desert riparian tree species

We investigated leaf gas exchange responses to leaf temperature, leaf-to-air vapor pressure deficit (VPD), and predawn and midday shoot water potential (psipd and psimd, respectively) of two native Sonoran Desert riparian tree species, Fremont cottonwood (Populus fremontii S. Wats.) and Goodding willow (Salix gooddingii Ball), and one exotic riparian tree species, saltcedar (Tamarix chinensis Lour. and related species). Measurements were made at two sites over 2 years that differed climatically. Because multiple linear regression models explained less than 29% of the variation in stomatal conductance (gs) and less than 48% of the variation in net photosynthetic rate (Pn) of all species, we used boundary-line analysis to compare gas exchange responses among species. Gas exchange rates were high in all species. The hyperbolic relationship between Pn and gs suggested that initial reductions in gs at high gs did not inhibit Pn. Reductions in gs of cottonwood and willow occurred at psimd values at or below previously reported xylem cavitation thresholds (-1.6 and -1.4 MPa, respectively), indicating tight stomatal regulation of water loss and a narrow cavitation safety margin. In contrast, reductions in gs of saltcedar occurred at psimd values well above the cavitation threshold (-7.0 MPa), but at much lower psimd values than in cottonwood and willow, suggesting a wider cavitation safety margin and less tight regulation of water loss in saltcedar. High VPD had a smaller effect on leaf gas exchange in willow than in cottonwood. In contrast, willow had a less negative psipd threshold for stomatal closure than cottonwood. Compared with cottonwood and willow, leaf gas exchange of saltcedar was more tolerant of high VPD and low psipd. These physiological characteristics of saltcedar explain its widespread success as an invader of riparian ecosystems containing native Fremont cottonwood and Goodding willow in the Sonoran Desert.     

Drought resistance of Ailanthus altissima: root hydraulics and water relations

Drought resistance of Ailanthus altissima (Mill.) Swingle is a major factor underlying the impressively wide expansion of this species in Europe and North America. We studied the specific mechanism used by A. altissima to withstand drought by subjecting potted seedlings to four irrigation regimes. At the end of the 13-week treatment period, soil water potential was -0.05 MPa for well-watered control seedlings (W) and -0.4, -0.8 and -1.7 MPa for drought-stressed seedlings (S) in irrigation regimes S1, S2 and S3, respectively. Root and shoot biomass production did not differ significantly among the four groups. A progressively marked stomatal closure was observed in drought-stressed seedlings, leading to homeostasis of leaf water potential, which was maintained well above the turgor loss point. Root and shoot hydraulics were measured with a high-pressure flow meter. When scaled by leaf surface area, shoot hydraulic conductance did not differ among the treated seedlings, whereas root hydraulic conductance decreased by about 20% in S1 and S2 seedlings and by about 70% in S3 seedlings, with respect to the well-watered control value. Similar differences were observed when root hydraulic conductance was scaled by root surface area, suggesting that roots had become less permeable to water. Anatomical observations of root cross sections revealed that S3 seedlings had shrunken cortical cells and a multilayer endodermal-like tissue that probably impaired soil-to-root stele water transport. We conclude that A. altissima seedlings are able to withstand drought by employing a highly effective water-saving mechanism that involves reduced water loss by leaves and reduced root hydraulic conductance. This water-saving mechanism helps explain how A. altissima successfully competes with native vegetation.     

Changes in root growth and relationships between plant organs and root hydraulic traits in American elm (Ulmus americana L.) and red oak (Quercus rubra L.) seedlings due to elevated CO2 level

The relationships between plant organs and root hydrological traits are not well known and the question arises whether elevated CO2 changes these relationships. This study attempted to answer this question. A pseudo-replicated experiment was conducted with two times 24 American elm (Ulmus americana L.) and 23 and 24 red oak (Quercus rubra L.) seedlings growing in ambient CO2 (around 360 μmol·L–1) and 540 ± 7.95 μmol·L–1 CO₂ in a greenhouse. After 71 days of treatment for American elm and 77 days for red oak, 14 American elm and 12 red oak seedlings from each of the two CO2 levels were randomly selected in order to examine the flow rate of root xylem sap, root hydraulic conductance, total root hydraulic conductivity, fine root and coarse root hydraulic conductivity. All seedlings were harvested to investigate total plant biomass, stem biomass and leaf biomass, leaf area, height, basal diameter, total root biomass, coarse root biomass and fine root biomass. The following conclusions are reached: 1) plant organs respond to the elevated CO2 level earlier than hydraulic traits of roots and may gradually lead to changes in hydraulic traits; 2) plant organs have different relationships with hydraulic traits of roots and elevated CO2 changes these relationships; the changes may be of importance for plants as means to acclimatize to changing environments; 3) biomass of coarse roots increased rather more than that of fine roots; 4) Lorentzian and Caussian models are better in estimating the biomass of seedlings than single-variable models.     

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.     

Changes in composition, structure and aboveground biomass over seventy-six years (1930-2006) in the Black Rock Forest, Hudson Highlands, southeastern New York State

We sought to quantify changes in tree species composition, forest structure and aboveground forest biomass (AGB) over 76 years (1930-2006) in the deciduous Black Rock Forest in southeastern New York, USA. We used data from periodic forest inventories, published floras and a set of eight long-term plots, along with species-specific allometric equations to estimate AGB and carbon content. Between the early 1930s and 2000, three species were extirpated from the forest (American elm (Ulmus americana L.), paper birch (Betula papyrifera Marsh.) and black spruce (Picea mariana (nigra) (Mill.) BSP)) and seven species invaded the forest (non-natives tree-of-heaven (Ailanthus altissima (Mill.) Swingle) and white poplar (Populus alba L.) and native, generally southerly distributed, southern catalpa (Catalpa bignonioides Walt.), cockspur hawthorn (Crataegus crus-galli L.), red mulberry (Morus rubra L.), eastern cottonwood (Populus deltoides Bartr.) and slippery elm (Ulmus rubra Muhl.)). Forest canopy was dominated by red oak and chestnut oak, but the understory tree community changed substantially from mixed oak-maple to red maple-black birch. Density decreased from an average of 1500 to 735 trees ha(-1), whereas basal area doubled from less than 15 m(2) ha(-1) to almost 30 m(2) ha(-1) by 2000. Forest-wide mean AGB from inventory data increased from about 71 Mg ha(-1) in 1930 to about 145 Mg ha(-1) in 1985, and mean AGB on the long-term plots increased from 75 Mg ha(-1) in 1936 to 218 Mg ha(-1) in 1998. Over 76 years, red oak (Quercus rubra L.) canopy trees stored carbon at about twice the rate of similar-sized canopy trees of other species. However, there has been a significant loss of live tree biomass as a result of canopy tree mortality since 1999. Important constraints on long-term biomass increment have included insect outbreaks and droughts.     

Carbon partitioning and allocation in northern red oak seedlings and mature trees in response to ozone

Northern red oak (Quercus rubra L.) seedlings and trees differ in their response to ozone. Previous work reported reductions in net photosynthesis, carboxylation efficiency and quantum yield of mature tree leaves, whereas seedling processes were unaffected by the same ozone exposure. To further characterize differences in ozone response between seedlings and mature trees, we examined carbon partitioning and allocation in 32-year-old trees and 4-year-old seedlings of northern red oak after exposure to subambient (seasonal SUM00 dose (sum of all hourly ozone exposures) = 31 ppm-h), ambient (SUM00 dose = 85 ppm-h) and twice ambient (SUM00 dose = 151 ppm-h) ozone concentrations for three growing seasons. For mature trees, ozone exposure decreased foliar starch partitioning, increased starch partitioning in branches and increased (14)C retention in leaves. In contrast, starch partitioning in leaves and branches, and foliar (14)C retention in seedlings were unaffected by ozone exposure, but soluble carbohydrate concentrations in coarse and fine roots of seedlings were reduced. Differences in carbohydrate demand between seedlings and mature trees may underlie the higher leaf ozone uptake rates and greater physiological response to ozone in mature northern red oak trees compared with seedlings.     

Drought effects on hydraulic conductivity and xylem vulnerability to embolism in diverse species and provenances of Mediterranean cedars

We studied hydraulic traits of young plants of the Mediterranean cedar species Cedrus atlantica (Endl.) G. Manetti ex Carrière (Luberon, France), C. brevifolia (Hook. f.) Henry (Cyprus), C. libani A. Rich (Hadeth El Jebbe, Lebanon) and C. libani (Armut Alani, Turkey). With an optimum water supply, no major differences were observed among species or provenances in either stem hydraulic conductivity (Ks) or leaf specific conductivity (Kl) measured on the main shoot. A moderate soil drought applied for 10 weeks induced marked acclimation through a reduction in Ks, particularly in the Lebanese provenance of C. libani, and a decrease in tracheid lumen size in all species. Cedrus atlantica, which had the smallest tracheids, was the species most vulnerable to embolism: a 50% loss in hydraulic conductivity (PsiPLC50) occurred at a water potential of -4.4 MPa in the well-watered treatment, and at -6.0 MPa in the moderate drought treatment. In the other species, PsiPLC50 was unaffected by moderate soil drought, and only declined sharply at water potentials between -6.4 and -7.5 MPa in both irrigation treatments. During severe drought, Ks of twigs and stomatal conductance (g(s)) were measured simultaneously as leaf water potential declined. For all species, lower vulnerability to embolism based on loss of Ks was recorded on current-year twigs. The threshold for stomatal closure (10% of maximum g(s)) was reached at a predawn water potential (Psi(pd)) of -2.5 MPa in C. atlantica (Luberon) and at -3.1 MPa in C. libani (Lebanon), whereas the other provenance and species had intermediate Psi(pd) values. Cedrus brevifolia, with a Psi(pd) (-3.0 MPa) close to that of C. libani (Lebanon), had the highest stomatal conductance of the study species. The importance of a margin of safety between water potential causing stomatal closure and that causing xylem embolism induction is discussed.     

Xylem conductivity and vulnerability to cavitation of ponderosa pine growing in contrasting climates

We examined the effects of increased transpiration demand on xylem hydraulic conductivity and vulnerability to cavitation of mature ponderosa pine (Pinus ponderosa Laws.) by comparing trees growing in contrasting climates. Previous studies determined that trees growing in warm and dry sites (desert) had half the leaf/sapwood area ratio (A(L)/A(S)) and more than twice the transpiration rate of trees growing in cool and moist sites (montane). We predicted that high transpiration rates would be associated with increased specific hydraulic conductivity (K(S)) and increased resistance to xylem cavitation. Desert trees had 19% higher K(S) than montane trees, primarily because of larger tracheid lumen diameters. Predawn water potential and water potential differences between the soil and the shoot were similar for desert and montane trees, suggesting that differences in tracheid anatomy, and therefore K(S), were caused primarily by temperature and evaporative demand, rather than soil drought. Vulnerability to xylem cavitation did not differ between desert and montane populations. A 50% loss in hydraulic conductivity occurred at water potentials between -2.61 and -2.65 MPa, and vulnerability to xylem cavitation did not vary with stem size. Minimum xylem tensions of desert and montane trees did not drop below -2.05 MPa. Foliage turgor loss point did not differ between climate groups and corresponded to mean minimum xylem tensions in the field. In addition to low A(L)/A(S), high K(S) in desert trees may provide a way to increase tree hydraulic conductivity in response to high evaporative demand and prevent xylem tensions from reaching values that cause catastrophic cavitation. In ponderosa pine, the flexible responses of A(L)/A(S) and K(S) to climate may preclude the existence of significant intraspecific variation in the vulnerability of xylem to cavitation.     

Survival and growth of upland oak and co-occurring competitor seedlings following single and repeated prescribed fires

Studies within and outside the U.S. indicate recurring oak (Quercus spp.) regeneration problems. In deciduous forests of the eastern U.S., a prevailing explanation for this trend is fire suppression leading to high competitor abundance and low understory light. In response, prescribed fire is increasingly used as a management tool to remedy these conditions and encourage future oak establishment and growth. Within eastern Kentucky, we implemented single and repeated (3×) prescribed fires over a 6-yr period (2002–2007). Pre- and post-burn, we quantified canopy cover and oak seedling survival and growth compared to other woody seedlings deemed potential competitors, primarily red maple (Acer rubrum L.) and sassafras (Sassafras albidum (Nutt.) Nees.). Burning temporarily decreased canopy cover 3–10%, but cover rebounded the subsequent growing season. Repeated burning ultimately produced canopy cover about 6% lower than sites unburned and burned once, suggesting a cumulative effect on understory light. Red maple exhibited low survival (∼40%) following single and repeated burns, but growth remained similar to unburned seedlings. Burning had little impact on sassafras survival and led to total height and basal diameters 2× greater than unburned seedlings. A single burn had no impact on red oak (Erythrobalanus spp.) survival and increased height and basal diameters 25–30%, but this positive growth response was driven by seedlings on several plots which experienced high burn temperatures and consequently high overstory mortality. White oaks (Leucobalanus spp.), however, exhibited twice as high mortality compared to those unburned, with no change in growth parameters. Repeated burning negatively impacted survival and growth of both oak groups compared to unburned seedlings. With both burn regimes, oaks with smaller pre-burn basal diameters exhibited the lowest post-burn survival. Thus, despite the ability of prescribed burns to temporarily increase understory light and reduce red maple survival, neither single or repeated burns placed oaks in an improved competitive position. These findings result from a combination of highly variable yet interdependent factors including the (1) life history traits of oaks compared to their co-occurring competitors, (2) pre-burn stature of pre-existing oak seedlings, and (3) variability in fire temperature and effects on understory light.