Hydraulic traits are associated with the distribution range of two closely related Mediterranean firs, Abies alba Mill. and Abies pinsapo Boiss

Abies alba and Abies pinsapo are two closely related fir species that occur in the Iberian Peninsula under very different environmental conditions. Abies alba proliferates in the humid European mountains, including the Spanish Pyrenees. In contrast, A. pinsapo is a relict species that occurs in some restricted areas of the Mediterranean mountain ranges in Spain and Morocco, which experience intense summer drought periods. To cope with the high atmospheric evaporative demand during summer, A. pinsapo may either have a high resistance to xylem cavitation or develop a very efficient conducting system to reduce the soil-to-leaf water potential gradient. To investigate such hypotheses, we measured (i) the xylem vulnerability to cavitation for different populations, and (ii) several anatomical and hydraulic parameters indicating xylem sufficiency for -supplying water to the shoot in two contrasting populations of both species. Our results show that the resistance to cavitation was not different between species or populations. However, hydraulic conductivity (K(h)), specific hydraulic conductivity (K(s)), leaf-specific conductivity (LSC) and whole-shoot hydraulic conductance (K(shoot)) were higher in A. pinsapo, indicating a higher efficiency of water transport, which should contribute to maintaining its xylem tension below the threshold for rapidly increasing cavitation. The higher K(s) in A. pinsapo was largely a result of its wider tracheids, suggesting that this species may be much more vulnerable to freeze-thaw-induced cavitation than A. alba. This is consistent with the absence of A. pinsapo in northern mountain ranges with cooler winters. These physiological differences could partly explain the niche segregation and the geographical separation of these two firs.     

Bole girdling affects metabolic properties and root, trunk and branch hydraulics of young ponderosa pine trees

Effects of trunk girdling on seasonal patterns of xylem water status, water transport and woody tissue metabolic properties were investigated in ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) trees. At the onset of summer, there was a sharp decrease in stomatal conductance (g(s)) in girdled trees followed by a full recovery after the first major rainfall in September. Eliminating the root as a carbohydrate sink by girdling induced a rapid reversible reduction in g(s). Respiratory potential (a laboratory measure of tissue-level respiration) increased above the girdle (branches and upper trunk) and decreased below the girdle (lower trunk and roots) relative to control trees during the growing season, but the effect was reversed after the first major rainfall. The increase in branch respiratory potential induced by girdling suggests that the decrease in g(s) was caused by the accumulation of carbohydrates above the girdle, which is consistent with an observed increase in leaf mass per area in the girdled trees. Trunk girdling did not affect native xylem embolism or xylem conductivity. Both treated and control trunks experienced loss of xylem conductivity ranging from 10% in spring to 30% in summer. Girdling reduced xylem growth and sapwood to leaf area ratio, which in turn reduced branch leaf specific conductivity (LSC). The girdling-induced reductions in g(s) and transpiration were associated with a decrease in leaf hydraulic conductance. Two years after girdling, when root-to-shoot phloem continuity had been restored, girdled trees had a reduced density of new wood, which increased xylem conductivity and whole-tree LSC, but also vulnerability to embolism.     

Patterns of hydraulic architecture and water relations of two tropical canopy trees with contrasting leaf phenologies: Ochroma pyramidale and Pseudobombax septenatum

Many authors have attempted to explain the adaptive response of tropical plants to drought based on studies of water relations at the leaf level. Little attention has been given to the role of the xylem system in the control of plant water requirements. To evaluate this role, we studied the hydraulic architecture and water relations parameters of two tropical canopy trees with contrasting leaf phenologies: deciduous Pseudobombax septenatum (Jacq.) Dug and evergreen Ochroma pyramidale (Cav. ex lamb) Urban, both in the family Bombacaceae. The hydraulic architecture parameters studied include hydraulic conductivity, specific conductivity, leaf specific conductivity, and Huber value. Water relations parameters include leaf water potential, stem and leaf water storage capacitance, transpiration, stomatal conductance, and vulnerability of stems to cavitation and loss of hydraulic conductivity by embolisms. Compared to temperate trees, both species showed a pattern of highly vulnerable stems (50% loss of conductivity due to embolism at water potentials less than 1 MPa) with high leaf specific conductivities. The vulnerability of xylem to water-stress-induced embolism was remarkably similar for the two species but the leaf specific conductivity of petioles and leaf-bearing stems of the evergreen species, Ochroma (e.g., 9.08 and 11.4 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively), were 3.4 and 2.3 times higher, respectively, than those of the deciduous species, Pseudobombax (e.g., 2.64 and 5.15 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively). A runaway embolism model was used to test the ability of Ochroma and Pseudobombax stems to maintain elevated transpiration rates during the higher evaporative demand of the dry season. The percent loss of leaf area predicted by the runaway embolism model for stems of Pseudobombax ranged from 5 to 30%, not enough to explain the deciduous phenology of this tree species without analysis of root resistance or leaf and petiole vulnerability to embolism.     

Regulation of water loss in populations of Populus trichocarpa: the role of stomatal control in preventing xylem cavitation

Variations in resistance to drought-induced xylem cavitation, xylem air-entry points, stomatal behavior, and hydraulic conductivity were measured in four populations of Populus trichocarpa Torr. & A. Gray collected along an east-west humidity and temperature gradient in Washington State, USA. Xylem air-entry points were less negative in trees from moist environments (-0.71 and -1.32 MPa in the Hoh and Nisqually populations, respectively) than in trees from dry environments (-1.55 and -1.67 MPa in the Palouse and Yakima populations, respectively). Xylem cavitation in response to experimental drought was consistent with air-injection measures of xylem air-entry points for a given population. Populations vulnerable to cavitation also exhibited higher stem specific hydraulic conductivities and limited stomatal control compared with resistant populations. Populations exhibiting vulnerability to cavitation and limited stomatal control desiccated more rapidly during drought compared with resistant populations. This study provides evidence of interpopulation variation in resistance to drought-induced xylem cavitation, stomatal behavior, and hydraulic conductivity within Populus trichocarpa.     

Hydraulic conductivity in roots of ponderosa pine infected with black-stain (Leptographium wageneri) or annosus (Heterobasidion annosum) root disease

Roots from healthy and diseased mature ponderosa pine, Pinus ponderosa Laws., trees were excavated from a site near Burns, Oregon. The diseased trees were infected with black-stain root disease, Leptographium wageneri Kendrick, or annosus root disease, Heterobasidion annosum (Fr.) Bref., or both. Axial hydraulic conductivity of the roots was measured under a positive head pressure of 5 kPa, and the conducting area was stained with safranin dye to determine specific conductivity (k(s)). In diseased roots, only 8-12% of the cross-sectional xylem area conducted water. Resin-soaked xylem completely restricted water transport and accounted for 13-16% of the loss in conducting area. In roots with black-stain root disease, 17% of the loss in conducting area was associated with unstained xylem, possibly resulting from occlusions or embolisms. Based on the entire cross-sectional area of infected roots, the k(s) of roots infected with black-stain root disease was 4.6% of that for healthy roots, whereas the k(s) of roots infected with annosus root disease was 2.6% of that for healthy roots. Although these low values were partly the result of the presence of a large number of diseased roots (72%) with no conducting xylem, the k(s) of functional xylem of diseased roots was only 33% of that for healthy roots. The low k(s) values of functional xylem in diseased roots may be caused by fungus induced occlusions preceding cavitation and embolism of tracheids. The k(s) of disease-free roots from diseased trees was only 70% of that for healthy roots from healthy trees. The disease-free roots had the same mean tracheid diameter and tissue density as the healthy roots, suggesting that the lower k(s) in disease-free roots of diseased trees may also have been caused by partial xylary occlusions.     

A case-study of water transport in co-occurring ring- versus diffuse-porous trees: contrasts in water-status, conducting capacity, cavitation and vessel refilling

Recent work has suggested that the large earlywood vessels of ring-porous trees can be extraordinarily vulnerable to cavitation making it necessary that these trees maintain a consistent and favorable water status. We compared cavitation resistance, vessel refilling, transport capacity and water status in a study of ring-porous Quercus gambelii Nutt. (oak) and diffuse-porous Acer grandidentatum Nutt. (maple). These species co-dominate summer-dry foothills in the western Rocky Mountains of the USA. Native embolism measurements, dye perfusions and balance pressure exudation patterns indicated that the large earlywood vessels of 2-3-year-old oak stems cavitated extensively on a daily basis as predicted from laboratory vulnerability curves, resulting in a more than 80% reduction in hydraulic conductivity. Maple branches showed virtually no cavitation. Oak vessels refilled on a daily basis, despite negative xylem pressure in the transpiration stream, indicating active pressurization of embo-lized vessels. Conductivity and whole-tree water use in oak were between about one-half and two-thirds that in maple on a stem-area basis; but were similar or greater on a leaf-area basis. Oak maintained steady and modest negative xylem pressure potentials during the growing season despite little rainfall, indicating isohydric water status and reliance on deep soil water. Maple was markedly anisohydric and developed more negative pressure potentials during drought, suggesting use of shallower soil water. Although ring porosity may have evolved as a mechanism for coping with winter freezing, this study suggests that it also has major consequences for xylem function during the growing season.     

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).     

Variation in drought resistance, drought acclimation and water conservation in four willow cultivars used for biomass production

Growth and water-use parameters of four willow (Salix spp.) clones grown in a moderate drought regime or with ample water supply were determined to characterize their water-use efficiency, drought resistance and capacity for drought acclimation. At the end of the 10-week, outdoor pot experiment, clonal differences were observed in: (1) water-use efficiency of aboveground biomass production (WUE); (2) resistance to xylem cavitation; and (3) stomatal conductance to leaf-specific, whole-plant hydraulic conductance ratio (g(st)/K(P); an indicator of water balance). Across clones and regimes, WUE was positively correlated with the assimilation rate to stomatal conductance ratio (A/g(st)), a measure of instantaneous water-use efficiency. Both of these water-use efficiency indicators were generally higher in drought-treated trees compared with well-watered trees. However, the between-treatment differences in (shoot-based) WUE were smaller than expected, considering the differences in A/g(st) for two of the clones, possibly because plants reallocated dry mass from shoots to roots when subject to drought. Higher root hydraulic conductance to shoot hydraulic conductance ratios (K(R)/K(S)) during drought supports this hypothesis. The same clones were also the most sensitive to xylem cavitation and, accordingly, showed the strongest reduction in g(st)/K(P) in response to drought. Drought acclimation was manifested in decreased g(st), g(st)/K(P), osmotic potential and leaf area to vessel internal cross-sectional area ratio, and increased K(R), K(P) and WUE. Increased resistance to stem xylem cavitation in response to drought was observed in only one clone. It is concluded that WUE and drought resistance traits are inter-linked and that both may be enhanced by selection and breeding.     

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.     

Whole-plant hydraulic resistance and vulnerability segmentation in Acer saccharinum

Hydraulic properties were studied in Acer saccharinum L., a riparian species that also grows well on a dry soil when transplanted. Hydraulic resistances were measured by two independent techniques: a new high-pressure flowmeter (HPFM) method and a conventional evaporative flux (EF) method. Vulnerability to cavitation was also investigated on petioles, stems and roots using a hydraulic conductivity technique. Vulnerability segmentation was found, i.e., roots, stems and petioles had different vulnerabilities to xylem dysfunction. Petioles were most vulnerable with 50% loss of hydraulic conductivity at -0.5 MPa, roots were least vulnerable (50% loss at -2.2 MPa) and stems were intermediate in vulnerability. The HPFM and the EF methods gave comparable results, except that the EF method gave a significantly higher value for resistance across petioles plus leaves. Native embolism was high enough to explain the discrepancy in resistance across petioles plus leaves between the HPFM and the EF methods, indicating that the HPFM estimates the minimum (potential) hydraulic resistance of plants. Whole-plant hydraulic resistance of A. saccharinum was low compared to resistances of other temperate species. The hydraulic characteristics of A. saccharinum were consistent with adaptation to its typical environment: low whole-plant resistance assures high transpiration rates in the presence of sufficient water, and vulnerability segmentation provides the ability to survive during droughts through shedding of expendable organs.     

Xylem vulnerability to cavitation in Pseudotsuga menziesii and Pinus ponderosa from contrasting habitats

In the Rocky Mountains, ponderosa pine (Pinus ponderosa (ssp.) ponderosa Dougl. ex P. Laws. & C. Laws) often co-occurs with Douglas-fir (Pseudotsuga menziesii var. glauca (Mayr) Franco). Despite previous reports showing higher shoot vulnerability to water-stress-induced cavitation in ponderosa pine, this species extends into drier habitats than Douglas-fir. We examined: (1) whether roots and shoots of ponderosa pine in riparian and slope habitats are more vulnerable to water-stress-induced cavitation than those of Douglas-fir; (2) whether species-specific differences in vulnerability translate into differences in specific conductivity in the field; and (3) whether the ability of ponderosa pine to extend into drier sites is a result of (a) greater plasticity in hydraulic properties or (b) functional or structural adjustments. Roots and shoots of ponderosa pine were significantly more vulnerable to water-stress-induced cavitation (overall mean cavitation pressure, Psi(50%) +/- SE = -3.11 +/- 0.32 MPa for shoots and -0.99 +/- 0.16 MPa for roots) than those of Douglas-fir (Psi(50%) +/- SE = -4.83 +/- 0.40 MPa for shoots and -2.12 +/- 0.35 MPa for roots). However, shoot specific conductivity did not differ between species in the field. For both species, roots were more vulnerable to cavitation than shoots. Overall, changes in vulnerability from riparian to slope habitats were small for both species. Greater declines in stomatal conductance as the summer proceeded, combined with higher allocation to sapwood and greater sapwood water storage, appeared to contribute to the ability of ponderosa pine to thrive in dry habitats despite relatively high vulnerability to water-stress-induced cavitation.     

How timing of stem girdling affects needle xylem structure in Scots pine

While needles represent a proportionally large fraction of whole-plant hydraulic resistance, no studies to date have investigated how source–sink disturbances affect needle xylem structure. In this study, we evaluated structural changes in xylem in current-year needles of Scots pine 227 and 411 days after stem girdling (hereafter referred to as DAG). Maximum and minimum tracheid lumen diameters and therefore also the size of tracheid lumen areas increased in needles 227 DAG compared to control needles. In contrast, tracheid dimensions were similar in needles 411 DAG as in the control needles, but smaller xylem area and lower number of tracheids resulted in the lower theoretical needle hydraulic conductivity of those needles. Several needle xylem parameters were intercorrelated in both control and girdled trees. These observed changes provide a new understanding of the processes that occur following a source–sink disturbance. Considering anatomical parameters such as the number of tracheids, tracheid dimension, or needle xylem area, which are rarely described in physiological studies, could be helpful, for example, in understanding to tree hydraulic systems or for modeling gas exchange. Finally, empirical equations were developed to calculate needle theoretical hydraulic conductivity and the number of tracheids in needles using an easily measurable parameter of needle xylem area.     

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.     

Linking hydraulic conductivity and photosynthesis to water-source partitioning in trees versus seedlings

The relationships between hydraulic and photosynthetic properties in plants have been widely studied, but much less is known about how these properties are linked to water-source partitioning, the spatial and temporal separation of water sources in ecosystems. Plant water-source partitioning is often influenced by the proximity of groundwater from the natural surface. We studied the water acquisition strategy and hydraulic and photosynthetic properties of Tuart (Eucalyptus gomphocephala D.C.), a large coastal tree species that occupies seasonally dry habitats underlain by superficial aquifers. Our goal was to quantify water-source partitioning as the proportion of xylem water derived from the vadose and saturated zones with respect to stage of development and proximity of groundwater. We then sought to associate the proportional contribution of a given water source with xylem hydraulic and photosynthetic properties, thus conferring a linkage. Seedlings were more inclined to use surface soil water when rainfall recharge of the upper profile occurred, suggesting that they maintained or rapidly developed a proportionally high amount of functional roots in the upper, seasonally dry, soil profile. This strategy was associated with a lower xylem-area-specific hydraulic conductivity (K(S)), leaf-area-specific hydraulic conductivity (K(L)) and maximum photon yield of photosystem II (F(V)/F(M)). In contrast, trees acquired water from a variety of sources in different seasons and had a higher K(S), K(L) and F(V)/F(M). Despite the higher K(S) and K(L) in trees, the midday hydrodynamic water potential gradient from soil to leaves, ΔΨ, was similar. We conclude that there was a linkage between hydraulic and photosynthetic properties with the partitioning of water sources and that this adaptation to long-term hydrological regimes accommodated the different hydraulic characteristics and hydrological environments of trees versus seedlings.     

Vulnerability of several conifers to air embolism

Hydraulic properties of xylem in seven species of conifer were studied during late winter and early spring 1991. Vulnerability to cavitation and air embolism was investigated using hydraulic conductivity and acoustic techniques. Embolisms were induced in branches excised from mature trees by air-drying them in the laboratory. Both techniques gave comparable results indicating that they both assess the same phenomenon. Within a tree, vulnerability was related to the permeability of the xylem, the largest stems tended to cavitate before the smallest ones when water deficits developed in a branch. Interspecific comparisons showed large differences in the xylem water potential needed to induce significant embolism, values ranged from -2.5 MPa in Pinus sylvestris to -4 MPa in Cedrus atlantica, but these differences did not correlate with differences in the xylem permeability of the species. The vulnerability of a species to air embolism was found to be consistent with its ecophysiological behavior in the presence of water stress, drought-tolerant species being less vulnerable than drought-avoiding species.     

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.     

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.     

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.     

Hydraulic and mechanical properties of young Norway spruce clones related to growth and wood structure

Stem segments of eight five-year-old Norway spruce (Picea abies (L.) Karst.) clones differing in growth characteristics were tested for maximum specific hydraulic conductivity (k(s100)), vulnerability to cavitation and behavior under mechanical stress. The vulnerability of the clones to cavitation was assessed by measuring the applied air pressure required to cause 12 and 50% loss of conductivity (Psi(12), Psi(50)) and the percent loss of conductivity at 4 MPa applied air pressure (PLC(4MPa)). The bending strength and stiffness and the axial compression strength and stiffness of the same stem segments were measured to characterize wood mechanical properties. Growth ring width, wood density, latewood percentage, lumen diameter, cell wall thickness, tracheid length and pit dimensions of earlywood cells, spiral grain and microfibril angles were examined to identify structure-function relationships. High k(s100) was strongly and positively related to spiral grain angle, which corresponded positively to tracheid length and pit dimensions. Spiral grain may reduce flow resistance of the bordered pits of the first earlywood tracheids, which are characterized by rounded tips and an equal distribution of pits along the entire length. Wood density was unrelated to hydraulic vulnerability parameters. Traits associated with higher hydraulic vulnerability were long tracheids, high latewood percentage and thick earlywood cell walls. The positive relationship between earlywood cell wall thickness and vulnerability to cavitation suggest that air seeding through the margo of bordered pits may occur in earlywood. There was a positive phenotypic and genotypic relationship between k(s100) and PLC(4MPa), and both parameters were positively related to tree growth rate. Variability in mechanical properties depended mostly on wood density, but also on the amount of compression wood. Accordingly, hydraulic conductivity and mechanical strength or stiffness showed no tradeoff.     

Water loss regulation in mature Hevea brasiliensis: effects of intermittent drought in the rainy season and hydraulic regulation

Effects of soil and atmospheric drought on whole-tree transpiration (E(T)), leaf water potential (Ψ(L)) and whole-tree hydraulic conductance (K(T)) were investigated in mature rubber trees (Hevea brasiliensis, clone RRIM 600) during the full canopy stage in the rainy season in a drought-prone area of northeast Thailand. Under well-watered soil conditions, transpiration was tightly regulated in response to high evaporative demand, i.e., above reference evapotranspiration (ET(0)) ~2.2 mm day(-1) or maximum vapor pressure deficit ~1.8 kPa. When the trees experienced intermittent soil drought E(T) decreased sharply when relative extractable water in the top soil was?相似文献