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.     

Coast live oak, Quercus agrifolia, susceptibility and response to goldspotted oak borer, Agrilus auroguttatus, injury in southern California

Oak mortality is often associated with a complex of decline factors. We describe the morphological and physiological responses of coast live oak, Quercus agrifolia Née, in California to an invasive insect, the goldspotted oak borer (GSOB), Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), and evaluate drought as a potential inciting factor. Morphological traits of 356 trees were assessed and physiological traits of 70 of these were monitored intensively over one growing season. Morphological characteristics of tree health included crown thinning and dieback; bole staining resulting from larval feeding; density of GSOB adult exit holes; and holes caused by woodpecker feeding. These characteristics were used to rank GSOB infestation/injury into four classes, and taken together, they explained 87% of the variation in a principal component analysis. Drought stress on various size/age and infestation classes of Q. agrifolia was measured by assessing branchlet pre-dawn and solar noon xylem water potential, leaf cell turgor potential, and water use efficiency over one growing season. Both morphological and physiological traits were highly variable in mature and old growth trees. Early summer plant water status (branchlet xylem water potential and water use efficiency) was similar between uninfested and newly colonized trees, suggesting that GSOB are not pre-selecting drought-stressed Q. agrifolia for oviposition. By late summer, leaf water and cell turgor potentials were lower in infested than in uninfested mature trees, suggesting that GSOB infestation causes drought stress in these trees. Among the tree size/age classes, infested old growth trees exhibited the greatest change in water use efficiency over the growing season, and showed greater morphological injury symptoms of decline than infested mature trees. Morphological attributes of decline in Q. agrifolia associated with GSOB were correlated weakly with increasing physiological drought stress among infestation classes of trees. We propose that the collection of morphological responses of Q. agrifolia to GSOB described here can be used to monitor the future expansion of the GSOB distribution as well as the GSOB-induced decline of Q. agrifolia in California.     

植物激素与木材形成

综述了植物激素在树木维管组织分化中的作用．生长素是调节木质部细胞分化的主要因子；细胞分裂素在诱导分生组织发端和分生细胞分化上起重要作用；赤霉素促进纤维的形成，赤霉素和生长素结合使用，可有效地促进次生木质部中纤维分化；乙烯能促进树木韧皮部和木质部分化。还讨沦了木材形成过程中的三个主要问题：生长素含量的多少直接控制了沿树轴木质部导管、管胞的大小和密度；外界环境因子影响木质部细胞分化；温带落叶阔叶树环孔材和散孔材形成的机理，生长素控制环孔材树种早材宽导管的形成以及晚材窄导管和纤维形成的假说。对今后改善树木木材的质和量上有一定参考价值。     

Differential summer water use by Pinus edulis and Juniperus osteosperma reflects contrasting hydraulic characteristics

Previous studies of pinyon-juniper woodlands show that Pinus edulis Engelm. makes better use of soil water from summer precipitation pulses than does co-occurring Juniperus osteosperma (Torr.) Little. To investigate the basis of this difference, we examined seasonal variation in cavitation and hydraulic conductance. Pinus edulis remained isohydric over the growing season. Minimum water potentials never fell below -2.3 MPa, and the extent of xylem cavitation remained near constant during the dry season. In contrast, J. osteosperma was anisohydric, reaching water potentials as low as -6.9 MPa, and experiencing progressively greater xylem cavitation as the dry season progressed despite having more cavitation-resistant xylem than P. edulis. We conducted an irrigation experiment to observe the responses of the study species to a summer pulse of water. Although sap flow increased in both species in response to the 25-mm irrigation pulse, only J. osteosperma responded to the 10-mm pulse. This was inconsistent with the response of P. edulis to light rain events and may have been due to a difference in the distribution of irrigation water and rain water between the under- and between-canopy areas. Whole-plant conductance increased following the 25-mm irrigation in P. edulis but remained constant in J. osteosperma. We hypothesized that this difference was caused, in part, by differential refilling of embolized xylem. Area specific hydraulic conductivity was 66% higher in roots of irrigated P. edulis trees relative to roots of control trees 3 days after the 25-mm irrigation (t = 2.14, P = 0.02, df = 16). There was no change in hydraulic conductivity of the roots of J. osteosperma or in the stems of either species. Our results indicate that the response to an irrigation pulse in P. edulis depended on cavitation avoidance in stems and the reversal of cavitation in roots, resulting in increased whole-plant conductance and water uptake. In contrast, J. osteosperma failed to exploit light summer rain events but was able to extract deep soil water at low water potentials.     

Drought tolerance,xylem sap abscisic acid and stomatal conductance during soil drying: a comparison of canopy trees of three temperate deciduous angiosperms

Patterns of water relations, xylem sap abscisic acid concentration ([ABA]) and stomatal aperture were characterized and compared in drought-sensitive black walnut (Juglans nigra L.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.) trees co-occurring in a second-growth forest in Missouri, USA. There were strong correlations among reduction in predawn leaf water potential, increased xylem sap [ABA] and stomatal closure in all species. Stomatal conductance was more closely correlated with xylem sap ABA concentration than with ABA flux or xylem sap pH and cation concentrations. In isohydric black walnut, increased concentrations of ABA in the xylem sap appeared to be primarily of root origin, causing stomatal closure in response to soil drying. In anisohydric sugar maple and white oak, however, there were reductions in midday leaf water potential associated with stomatal closure, making it uncertain whether drought-induced xylem sap ABA was of leaf or root origin. The role of root-originated xylem sap ABA in these species as a signal to the shoot of the water status of the roots is, therefore, less certain.     

Scaling of angiosperm xylem structure with safety and efficiency

We tested the hypothesis that greater cavitation resistance correlates with less total inter-vessel pit area per vessel (the pit area hypothesis) and evaluated a trade-off between cavitation safety and transport efficiency. Fourteen species of diverse growth form (vine, ring- and diffuse-porous tree, shrub) and family affinity were added to published data predominately from the Rosaceae (29 species total). Two types of vulnerability-to-cavitation curves were found. Ring-porous trees and vines showed an abrupt drop in hydraulic conductivity with increasing negative pressure, whereas hydraulic conductivity in diffuse-porous species generally decreased gradually. The ring-porous type curve was not an artifact of the centrifuge method because it was obtained also with the air-injection technique. A safety versus efficiency trade-off was evident when curves were compared across species: for a given pressure, there was a limited range of optimal vulnerability curves. The pit area hypothesis was supported by a strong relationship (r2 = 0.77) between increasing cavitation resistance and diminishing pit membrane area per vessel (A(P)). Small A(P) was associated with small vessel surface area and hence narrow vessel diameter (D) and short vessel length (L)--consistent with an increase in vessel flow resistance with cavitation resistance. This trade-off was amplified at the tissue level by an increase in xylem/vessel area ratio with cavitation resistance. Ring-porous species were more efficient than diffuse-porous species on a vessel basis but not on a xylem basis owing to higher xylem/vessel area ratios in ring-porous anatomy. Across four orders of magnitude, lumen and end-wall resistivities maintained a relatively tight proportionality with a near-optimal mean of 56% of the total vessel resistivity residing in the end-wall. This was consistent with an underlying scaling of L to D(3/2) across species. Pit flow resistance did not increase with cavitation safety, suggesting that cavitation pressure was not related to mean pit membrane porosity.     

Contrasting distribution and seasonal dynamics of carbohydrate reserves in stem wood of adult ring-porous sessile oak and diffuse-porous beech trees

We tested the hypothesis that broad-leaved forest species with contrasting wood anatomy and hydraulic system (ring-porous versus diffuse-porous) also differ in distribution and seasonal dynamics of carbohydrate reserves in stem wood. Total nonstructural carbohydrate (TNC) reserves (starch and sugars) were measured enzymatically in the 10 youngest stem xylem rings of adult oak (Quercus petraea (Matt.) Liebl.) and beech (Fagus sylvatica L.) trees during an annual cycle. Radial distribution of carbohydrates was investigated according to ring age. On all dates, oak trees had twofold higher TNC concentration than beech trees (41 versus 23 mg g(DM)(-1)), with starch accounting for the high TNC concentration in oak. Seasonal dynamics of TNC concentration were significantly (P < 0.05) more pronounced in oak (20-64 mg TNC g(DM)(-1)) than in beech (17-34 mg TNC g(DM)(-1)). A marked decrease in TNC concentration was observed in oak trees during bud burst and early wood growth, whereas seasonal fluctuations in TNC concentrations in beech trees were small. The radial distribution of TNC based on ring age differed between species: TNC was restricted to the sapwood rings in oak, whereas in beech, it was distributed throughout the wood from the outermost sapwood ring to the pith. Although the high TNC concentrations in the outermost rings accounted for most of the observed seasonal pattern, all of the 10 youngest xylem rings analyzed participated in the seasonal dynamics of TNC in beech trees. The innermost sapwood rings of oak trees had low TNC concentrations. Stem growth and accumulation of carbon reserves occurred concomitantly during the first part of the season, when there was no soil water deficit. When soil water content was depleted, stem growth ceased in both species, whereas TNC accumulation was negligibly affected and continued until leaf fall. The contrasting dynamics and distribution of carbohydrate reserves in oak and beech are discussed with reference to differences in phenology, early spring growth and hydraulic properties between ring-porous trees and diffuse-porous trees.     

Stem respiration in a closed-canopy upland oak forest

Stem respiration was measured throughout 1993 on 56 mature trees of three species (Quercus alba L., Quercus prinus L., and Acer rubrum L.) in Walker Branch Watershed, Oak Ridge, Tennessee. A subset of the trees was remeasured during 1994. Diameter increments, stem temperatures and soil water were also monitored. Respiration rates in the spring and summer of 1993 tracked growth rate increments, except during a drought when growth dropped to zero and respiration increased to its highest rate. During the dormant season, rates of total stem respiration (R(t)) tended to be greater in large trees with thick sapwood but no such trend was observed during the growing season. Before and after the growing season, respiration rates correlated well with stem temperatures. Estimated values of Q(10) were 2.4 for the two oak species and 1.7 for red maple. The Q(10) values were used along with baseline respiration measurements and stem temperatures to predict seasonal changes in maintenance respiration (R(m)). In red maple, annual total R(m) accounted for 56 and 60% of R(t) in 1993 and 1994, respectively. In chestnut oak, R(m) accounted for 65 and 58% of R(t) in 1993 and 1994, respectively. In white oak, R(m) accounted for 47 and 53% of R(t) in 1993 and 1994, respectively. Extrapolating these data to the stand level showed that woody tissue respiration accounted for 149 and 204 g C m(-2) soil surface year(-1) in 1993 and 1994, respectively.     

The anatomical traits of trunk wood and their relevance to oak (Quercus robur L.) vitality

Oaks’ decline in vitality is attributed to a complex process that involves interactions of several factors leading to increased trees’ mortality. This study investigates the structure of trunk wood of oaks with reference to its physiological role in hydraulic conductivity. On the basis of the crown condition, the oaks were classified into three health groups: healthy trees, declining trees and dead trees. Anatomical traits of wood, such as annual ring width, vessel density, vessel diameter of earlywood and theoretical hydraulic conductivity, were measured and calculated. The narrowest annual rings formed by the cambium were observed in dead oaks. These trees were also characterized by the smallest diameter of earlywood vessels, not only in the period of occurrence of dieback symptoms, but also during their whole life. It is suggested that the formation of narrow annual rings and earlywood vessels of small diameter increases susceptibility of a tree to decay. A reduced vessel diameter implies changes in hydraulic conductivity of oak trunks and thus impairs the water transport, which affects the health of trees. The process of oak decline is considered to have characteristics of natural selection and leads to the elimination of the weakest trees.     

Xylem hydraulic adjustment and growth response of Quercus canariensis Willd. to climatic variability

Global change challenges forest adaptability at the distributional limit of species. We studied ring-porous Quercus canariensis Willd. xylem traits to analyze how they adjust to spatio-temporal variability in climate. Trees were sampled along altitudinal transects, and annual time series of radial growth (ring width (RW)) and several earlywood vessel (EV) traits were built to analyze their relationships with climate. The trees responded to increasing water constraints with decreasing altitude and changes in climate in the short term but the analyses showed that xylem did not acclimate in response to long-term temperature increase during the past 30 years. The plants' adjustment to climate variability was expressed in a different but complementary manner by the different xylem traits. At low elevations, trees exhibited higher correlations with water stress indices and trees acclimated to more xeric conditions at low elevations by reducing radial growth and hydraulic diameter (D(H)) but increasing the density of vessels (DV). Average potential conductivity (K(H)) was similar for trees at different altitudes. However, inter-tree differences in xylem traits were higher than those between altitudes, suggesting a strong influence of individual genetic features or micro-site conditions. Trees exhibited higher RW those years with larger D(H) and particularly the linear density of vessels (DV(l)), but partly, climatic signals expressed in RW differed from those in EVs. Trees produced larger D(H) after cold winters and wet years. Ring width responded positively to wet and cool weather in fall and spring, whereas the response to climate of DV and K(H) was generally opposite to that of RW. These relationships likely expressed the negative impact of high respiration rates in winter on the carbon pools used to produce the EVs in the next spring and the overall positive influence of water availability for trees. Our results showed that trees at different sites were able to adjust their hydraulic architecture to climatic variability and temperature increase during recent decades coordinating several complementary traits. Nonetheless, it should be monitored whether they will succeed to acclimate to future climatic scenarios of increasing water stress.     

Drought tolerance,xylem sap abscisic acid and stomatal conductance during soil drying: a comparison of young plants of four temperate deciduous angiosperms

Patterns of water relations, xylem sap abscisic acid (ABA) concentration ([ABA]) and stomatal aperture were compared in drought-sensitive black walnut (Juglans nigra L.) and black willow (Salix nigra Marsh.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.). Strong correlations among reduction in predawn water potential, increase in xylem sap [ABA] and stomatal closure were observed in all species. Stomatal response was more highly correlated with xylem [ABA] than with ABA flux. Xylem sap pH and ion concentrations appeared not to play a major role in the stomatal response of these species. Stomata were more sensitive to relative changes in [ABA] in drought-sensitive black walnut and black willow than in sugar maple and white oak. In the early stages of drought, increased [ABA] in the xylem sap of black walnut and black willow was probably of root origin and provided a signal to the shoot of the water status of the roots. In sugar maple and white oak, leaf water potential declined with the onset of stomatal closure, so that stomatal closure also may have occurred in response to the change in leaf water potential.     

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.     

内聚力-张力学说的新证据(英文)

虽然内聚力 -张力学说已经建立了一个多世纪 ,但要完全接受它仍有一些问题 ,因为它的一个推论是很难证实的 ,即木质部导管中存在大的负压 .特别是近年来木质部压力探针和Z型管试验的结果表明 ,引起空穴的木质部压力仅在 - 0 5MPa以上 .有人根据这一结果认为必须对木质部水分运输的内聚力 -张力学说加以修正 .但是 ,目前又有两个研究小组应用相似改良的Z型管方法测定了产生空穴的木质部压力 .他们的结果显示 ,不同树种木质部导管保持水分传导的压力范围在 - 1 2MPa到- 3 5MPa以下 .这表明树木木质部导管中存在较大负压 ,从而有力支持了内聚力 -张力学说     

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 in roots and stems of Douglas-fir and white fir

Roots of hardwoods have been shown to be more vulnerable to xylem cavitation than stems. This study examined whether this pattern is also observed in a conifer species. Vulnerability to cavitation was determined from the pressure required to inject air into the vascular system of hydrated roots and stems, and reduce hydraulic conductance of the xylem. According to the air-seeding hypothesis for the cavitation mechanism, these air pressures predict the negative xylem pressure causing cavitation in dehydrating stems. This was evaluated for stems of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and white fir (Abies concolor (Gord. & Glend.) Lindl.). The air-injection method was applied to roots and stems of different sizes and positions in Douglas-fir trees. Roots, especially smaller roots with a xylem diameter < 5 mm, were more vulnerable to cavitation than stems. Mean cavitation pressure for smaller roots was -2.09 +/- 0.42 versus -3.80 +/- 0.19 MPa for larger roots (> 8 mm diameter). Within the shoot system, smaller stems (< 5 mm diameter) were most vulnerable to cavitation, having a mean cavitation pressure of -4.23 +/- 0.565 versus -5.27 +/- 0.513 MPa for large stems (> 8 mm diameter). There was no correlation between tracheid diameter and mean cavitation pressure within root or stem systems, despite larger tracheid diameters in roots (23.3 +/- 3.9 micro m) than in stems (9.2 +/- 1.6 micro m). Smaller safety margins from cavitation in roots may be beneficial in limiting water use during mild drought, and in protecting the stem from low xylem pressures during extreme drought.     

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.     

Associations between growth, wood anatomy, carbon isotope discrimination and mortality in a Quercus robur forest

Observations of forest mortality are increasing globally, but relatively little is known regarding the underlying mechanisms driving these events. Tree rings carry physiological signatures that may be used as a tool for retrospective analyses. We capitalized on a local soil water drainage event in 1982 that resulted in increased mortality within a stand of oak trees (Quercus robur), to examine the underlying physiological patterns associated with survival and death in response to soil water limitations. Pre-dawn water potentials showed more negative values for trees in the process of dying compared with those that survived. We used tree rings formed over the 123 years prior to mortality to estimate productivity from basal area increment (BAI, mm(2)), multiple xylem hydraulic parameters via anatomical measurements and crown-level gas exchange via carbon isotope discrimination (Δ, ‰). Oaks that died had significantly higher BAI values than trees that survived until the drainage event, after which the BAI of trees that died declined dramatically. Hydraulic diameter and conductivity of vessels in trees that died were higher than in surviving trees until the last 5 years prior to mortality, at which time both groups had similar values. Trees that died had consistently lower Δ values than trees that survived. Therefore, tree mortality in this stand was associated with physiological differences prior to the onset of soil water reduction. We propose that trees that died may have been hydraulically underbuilt for dry conditions, which predisposes them to severe hydraulic constraints and subsequent mortality. Measurements of above-ground/below-ground dry mass partitioning will be critical to future tests of this hypothesis. Based on these results, it is probable that pedunculate oak trees will experience greater future mortality if climate changes cause more severe droughts than the trees have experienced previously.     

Patterns of stomatal conductance among blue oak (Quercus douglasii) size classes and populations: implications for seedling establishment

Quercus douglasii Hook. & Arn. (blue oak) is a deciduous white oak that is currently failing to regenerate throughout much of its range in California, USA. Patterns of water use were observed in adult trees, saplings and seedlings to determine if ontogenetic changes in water use occur, which might be important in the establishment of this long-lived perennial species in a Mediterranean-type system. Seasonal and diurnal stomatal conductance (g(s)), late-season predawn xylem water potentials (Psi(pre)), carbon isotopic ratio (delta(13)C) and soil water status were compared among the three size classes at three sites differing in mean precipitation and soil water characteristics. Comparisons were also made between microsites with and without regeneration (defined by the presence or absence of saplings). Overall patterns of water use were consistent among the three sites, except that, at the site with the highest rainfall, Q. douglasii plants had higher g(s) and more positive Psi(pre) values. Although no differences in water use patterns were found between regeneration and non-regeneration microsites, the observed ontogenetic differences in water use may have important implications for Q. douglasii establishment. Compared with adult trees and saplings, seedlings had higher gas exchange rates during periods of high soil water content (early in the season and in the morning). Seedling g(s) was correlated with percent extractable soil water (ESW) throughout the season; adult tree and sapling g(s) was correlated with ESW between June and September. Despite experiencing greater water stress (indicated by more negative Psi(pre) values) than older trees, seedlings had more negative delta(13)C values, implying lower water-use efficiencies.     

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.     

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.