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.     

Calibration of thermal dissipation sap flow probes for ring- and diffuse-porous trees

Thermal dissipation probes (the Granier method) are routinely used in forest ecology and water balance studies to estimate whole-tree transpiration. This method utilizes an empirically derived equation to measure sap flux density, which has been reported as independent of wood characteristics. However, errors in calculated sap flux density may occur when large gradients in sap velocity occur along the sensor length or when sensors are inserted into non-conducting wood. These may be conditions routinely associated with ring-porous species, yet there are few cases in which the original calibration has been validated for ring-porous species. We report results from laboratory calibration measurements conducted on excised stems of four ring-porous species and two diffuse-porous species. Our calibration results for ring-porous species were considerably different compared with the original calibration equation. Calibration equation coefficients obtained in this study differed by as much as two to almost three orders of magnitude when compared with the original equation of Granier. Coefficients also differed between ring-porous species across all pressure gradient conditions considered; however, no differences between calibration slopes were observed for data collected within the range of expected in situ pressure gradients. In addition, dye perfusions showed that in three of the four ring-porous species considered, active sapwood was limited to the outermost growth ring. In contrast, our calibration results for diffuse-porous species showed generally good agreement with the empirically derived Granier calibration, and dye perfusions showed that active sapwood was associated with many annual growth rings. Our results suggest that the original calibration of Granier is not universally applicable to all species and xylem types and that previous estimates of absolute rates of water use for ring-porous species obtained using the original calibration coefficients may be associated with substantial error.     

Dynamic variation in sapwood specific conductivity in six woody species

Our goals were to quantify how non-embolism-inducing pressure gradients influence trunk sapwood specific conductivity (k(s)) and to compare the impacts of constant and varying pressure gradients on k(s) with KCl and H2O as the perfusion solutions. We studied six woody species (three conifers and three angiosperms) which varied in pit membrane structure, pit size and frequency of axial water transport across pits (long versus short conduits). Both stepwise ("steady") and nonlinear continuous ("non-steady") decreases in the pressure gradient led to decreased k(s) in all species but white oak (Quercus garryana Dougl. ex Hook), a ring-porous and long-vesseled angiosperm. In one diffuse-porous angiosperm (red alder, Alnus rubra Bong.) and two conifers (western red cedar, Thuja plicata Donn. ex D. Don, and Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco), k(s) was 10-30% higher under steady pressure gradients than under non-steady pressure gradients, and a decrease in the pressure gradient from 0.15 to 0.01 MPa m(-1) caused a 20-42% decrease in k(s). In another diffuse-porous angiosperm (maple, Acer macrophyllum Pursh) and in a third coniferous species (western hemlock, Tsuga heterophylla (Raf.) Sarg), there was no difference between k(s) measured under steady and non-steady pressure gradients. With the exception of western red cedar, a conifer with simple pit membranes, the differences in k(s) between low and high pressure gradients tended to be lower in the conifers than in the diffuse-porous angiosperms. In Douglas-fir, western red cedar and the diffuse-porous angiosperms, k(s) was higher when measured with KCl than with H2O. In white oak, there were no differences in k(s) whether measured under steady or non-steady pressure gradients, or when xylem was perfused with KCl or H2O. The species differences in the behavior of k(s) suggest that elasticity of the pit membrane was the main factor causing k(s) to be disproportionate to the pressure gradient and to the different pressure regimes. The results imply that, if nonlinearities in pressure-flux relationships are ignored when modeling tree water relations in vivo, large errors will result in the predictions of tree water status and its impact on stomatal control of transpiration and photosynthesis.     

Xylem vulnerability to cavitation varies among poplar and willow clones and correlates with yield

Xylem vulnerability to cavitation is a promising criterion for identifying trees with high drought tolerance, but traditional techniques for measuring cavitation resistance are unsuitable for screening large numbers of genotypes. We tested the potential of the new Cavitron technique for high throughput screening of cavitation resistance in five poplar (Populus spp.) and four willow (Salix spp.) clones. The Cavitron technique enabled the screening of three to four clones per day with sufficient accuracy to reveal significant differences between clones. Because intraspecific screening may be better carried out through the identification of correlated and more easily measured traits, we attempted to identify accessible parameters that correlate to cavitation resistance. Variability in vulnerability to cavitation across clones was poorly correlated with anatomical traits such as vessel diameter, vessel wall strength, wood density and fiber wall thickness; however, a striking correlation was established between cavitation resistance and aboveground biomass production, indicating a possible trade-off between xylem safety and growth potential.     

Shoot and root vulnerability to xylem cavitation in four populations of Douglas-fir seedlings

The objectives of this study were to assess the range of genotypic variation in the vulnerability of the shoot and root xylem of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings to water-stress-induced cavitation, and to assess the trade-off between vulnerability to cavitation and conductivity per unit of stem cross-sectional area (k(s)), both within a species and within an individual tree. Douglas-fir occupies a broad range of environments and exhibits considerable genetic variation for growth, morphology, and drought hardiness. We chose two populations from each of two varieties (the coastal var. menziesii and the interior var. glauca) to represent environmental extremes of the species. Vulnerability curves were constructed for shoots and roots by plotting the percentage loss in conductivity versus water potential. Vulnerability in shoot and root xylem varied genetically with source climate. Stem xylem differed in vulnerability to cavitation between populations; the most mesic population, coastal wet (CW), was the most susceptible of the four populations. In the roots, the most vulnerable population was again CW; the interior wet (IW) population was moderately susceptible compared with the two dry populations, coastal dry (CD) and interior dry (ID). Root xylem was more susceptible to cavitation than stem xylem and had significantly greater k(s). The trade-off between vulnerability to cavitation and k(s), however, was not evident across populations. The most vulnerable population (CW) had a shoot k(s) of 0.534 +/- 0.067 &mgr;mol m(-2) s(-1) MPa(-1), compared with 0.734 +/- 0.067 &mgr;mol m(-2) s(-1) MPa(-1) for the less vulnerable CD stems. In the roots, IW was more vulnerable than ID, but had the same k(s).     

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.     

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.     

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。     

Patterns in hydraulic architecture and their implications for transport efficiency

We evaluated whether patterns in hydraulic architecture increase transport efficiency. Five patterns are identified: area-preserving branching; variable trunk versus twig sap velocity; distally decreasing leaf specific conductivity (K(L)) and conduit diameter; and a decline in leaf specific conductance (k(L)) of the entire plant with maturation. These patterns coexist in innumerable combinations depending on the ratio of distal/proximal conduit number (F). The model of West and colleagues does not account for this diversity, in part by specifying F = 1 and requiring a specific conduit taper derived from the incorrect premise that k(L) is constant with plant size. We used Murray's law to identify the conduit taper that maximizes k(L)for a given vascular investment. Optimal taper requires the ratio of distal/proximal conduit diameter to equal the ratio of distal/proximal K(L). The smaller these ratios, the greater the k(L). Smaller ratios are achieved by an increase in F. Conductivity and diameter ratios < 1 and F >/= 1 in plants are therefore consistent with maximizing conducting efficiency. However, the benefit of increasing F requires area-increasing conduit branching, potentially leading to mechanical instability of trees. This trade-off may explain why tree stems were relatively inefficient with F near 1 and limited conduit taper compared with vine stems or compound leaves with F > 1 and greater taper. Within trees, the anatomies of a coniferous and a diffuse-porous species were less efficient than that of a ring-porous species, presumably because the latter allows conduit area to increase distally without also increasing total xylem area. This is consistent with decelerating sap velocities from trunk to twigs in ring-porous trees versus accelerating velocities in other types. In general, the observed architectural patterns are consistent with the maximization of transport efficiency operating within mechanical constraints.     

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.     

Physical analysis of the process of cavitation in xylem sap

Recent studies have confirmed that cavitation in xylem is caused by air bubbles. We analyzed expansion of a preexistent bubble adhering to a crack in a conduit wall and a bubble formed by the passage of air through a pore of a pit membrane, a process known as air seeding. We consider that there are two equilibrium states for a very small air bubble in the xylem: one is temporarily stable with a bubble radius r1 at point s1 on the curve P(r) relating pressure within the bubble (P) with bubble radius (r); the other is unstable with a bubble radius r2 at point s2 on Pr (where r1 < r2). In each equilibrium state, the bubble collapse pressure (2sigma/r, where sigma is surface tension of water) is balanced by the pressure difference across its surface. In the case of a bubble from a crack in a conduit wall, which is initially at point s1, expansion will occur steadily as water potential decreases. The bubble will burst only if the xylem pressure drops below a threshold value. A formula giving the threshold pressure for bubble bursting is proposed. In the case of an air seed entering a xylem conduit through a pore in a pit membrane, its initial radius may be r2 (i.e., the radius of the pore by which the air seed entered the vessel) at point s2 on Pr. Because the bubble is in an unstable equilibrium when entering the conduit, it can either expand or contract to point s1. As water vaporizes into the air bubble at s2, P rises until it exceeds the gas pressure that keeps the bubble in equilibrium, at which point the bubble will burst and induce a cavitation event in accordance with the air-seeding hypothesis. However, other possible perturbations could make the air-seeded bubble contract to s1, in which case the bubble will burst at a threshold pressure proposed for a bubble expanding from a crack in a conduit wall. For this reason some cavitation events may take place at a xylem threshold pressure (Pl'*) other than that determined by the formula, Plp'* = -2sigma/rp, proposed by Sperry and Tyree (1988), which is applicable only to air-seeded bubbles at s2. The more general formula we propose for calculating the threshold pressure for bubble breaking is consistent with the results of published experiments.     

用根压法研究竹子的耐旱、耐寒性

水分在植物体内长距离传输是植物水分研究中一个很重要的研究方向(Tyree,1997;Steudle,2001;Zimmermann et al.,2002;2004)。木质部管道分子(导管或管胞)是植物体内水分长距离运输     

The trade-off between safety and efficiency in hydraulic architecture in 31 woody species in a karst area

Karst topography is a special landscape shaped by the dissolution of one or more layers of soluble bedrock, usually carbonate rock such as limestone or dolomite. Due to subterranean drainage, overland flow, extraction of water by plants and evapotranspiration, there may be very limited surface water. The hydraulic architecture that plants use to adapt to karst topography is very interesting, but few systematic reports exist. The karst area in southwestern China is unique when compared with other karst areas at similar latitudes, because of its abundant precipitation, with rainfall concentrated in the growing season. In theory, resistance to water-stress-induced cavitation via air seeding should be accompanied by decreased pore hydraulic conductivity and stem hydraulic conductivity. However, evidence for such trade-offs across species is ambiguous. We measured the hydraulic structure and foliar stable carbon isotope ratios of 31 karst woody plants at three locations in Guizhou Province, China, to evaluate the functional coordination between resistance to cavitation and specific conductivity. We also applied phylogenetically independent contrast (PIC) analysis in situations where the inter-species correlations of functional traits may be biased on the potential similarity of closely related species. The average xylem tension measurement, at which 50% of hydraulic conductivity of the plants was lost (Ψ(50)), was only -1.27 MPa. Stem Ψ(50) was positively associated with specific conductance (K(s)) (P?相似文献   

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.     

A Discussion on Drought Tolerance Through Hydraulic Architecture of Woody Plants

In this paper, ten woody plants grew on the campus of the Beijing Forestry University were selected for measuring hydraulic architecture parameters and water potential of one-year-old twigs. The results show that day and night change of water potential and hydraulic architecture parameters appear to be obviously convex trend. The relationship models between water potential and hydraulic conductivity, special conductivity and leaf special conductivity were respectively established, which were simulated by equations Y=axb and Y=ax2 bx c, through which the parameters physiological meanings were discussed. The results show also show that embolism vulnerability of ten tree species was Robinia pseudoacacia> Gleditsia sinensis> Acer truncatum> Prunus davidiana> Cornus alba> Platycladus orientalis> Cedrus deodara> Pinus bungeana> Pinus tabulaeformis> Jasminum nudiflorum, and that the water transport efficiency in xylem and embolism vulnerability of the species are ring-porous species> diffuse-porous species>non     

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.     

Variation in seedling hydraulic architecture as a function of species and environment

Northern red oak (Quercus rubra L.) and yellow-poplar (Liriodendron tulipifera L.) were grown for two years in full sunlight (unshaded) or 20% of full sunlight (shaded) under either well-watered or drought conditions. There was a close association between evaporative flux (in situ) and leaf-specific conductivity (LSC) in stem segments of both species. Shaded, drought-stressed seedlings of both species had significantly reduced leaf area, evaporative flux, volume flow rate in xylem, flow velocity, potentially functional xylem area, and LSC than unshaded, well-watered seedlings. Reductions in LSC associated with drought or shade were similar for both species; and within a treatment, both species had similar LSC. Species differed in the manner of LSC adjustment to drought and shade. Reductions in leaf area associated with drought or shade were accompanied primarily by reductions in potentially functional xylem area for L. tulipifera, and by reductions in flow velocity for Q. rubra. These results suggest (1) the existence of a homeostatic balance between evaporative flux and LSC, (2) that species with widely different growth patterns and xylem anatomies may develop similar LSC within the same environment, and (3) a possible hydraulic basis for differences in habitat between ring- and diffuse-porous species.     

Transpiration sensitivity of urban trees in a semi-arid climate is constrained by xylem vulnerability to cavitation

Establishing quantitative links between plant hydraulic properties and the response of transpiration to environmental factors such as atmospheric vapor pressure deficit (D) is essential for improving our ability to understand plant water relations across a wide range of species and environmental conditions. We studied stomatal responses to D in irrigated trees in the urban landscape of Los Angeles, California. We found a strong linear relationship between the sensitivity of tree-level transpiration estimated from sap flux (m(T); slope of the relationship between tree transpiration and ln D) and transpiration at D=1 kPa (E(Tref)) that was similar to previous surveys of stomatal behavior in natural environments. In addition, m(T) was significantly related to vulnerability to cavitation of branches (P(50)). While m(T) did not appear to differ between ring- and diffuse-porous species, the relationship between m(T) and P(50) was distinct by wood anatomy. Therefore, our study confirms systematic differences in water relations in ring- versus diffuse-porous species, but these differences appear to be more strongly related to the relationship between stomatal sensitivity to D and vulnerability to cavitation rather than to stomatal sensitivity per se.     

Changes in petiole hydraulic properties and leaf water flow in birch and oak saplings in a CO2-enriched atmosphere

Water relations in woody species are intimately related to xylem hydraulic properties. High CO(2) concentrations ([CO(2)]) generally decrease transpiration and stomatal conductance (g(s)), but there is little information about the effect of atmospheric [CO(2)] on xylem hydraulic properties. To determine the relationship between water flow and hydraulic structure at high [CO(2)], we investigated responses of sun and shade leaves of 4-year-old saplings of diffuse-porous Betula maximowicziana Regel and ring-porous Quercus mongolica Fisch. ex Ledeb. ssp. crispula (Blume) Menitsky grown on fertile brown forest soil or infertile volcanic ash soil and exposed to 500 micromol CO(2) mol(-1) for 3 years. Regardless of species and soil type, elevated [CO(2)] consistently decreased water flow (i.e., g(s) and leaf-specific hydraulic conductivity) and total vessel area of the petiole in sun leaves; however, it had no effect on these parameters in shade leaves, perhaps because g(s) of shade leaves was already low. Changes in water flow at elevated [CO(2)] were associated with changes in petiole hydraulic properties.     

Interspecific variation in xylem vulnerability to cavitation among tropical tree and shrub species

In tropical moist forests, seasonal drought limits plant survival, productivity and diversity. Drought-tolerance mechanisms of tropical species should reflect the maximum seasonal water deficits experienced in a particular habitat. We investigated stem xylem vulnerability to cavitation in nine tropical species with different life histories and habitat associations. Stem xylem vulnerability was scored as the xylem water potential causing 50 and 75% loss of hydraulic conductivity (P50 and P75, respectively). Four shade-tolerant shrubs ranged from moderately resistant (P50=-1.9 MPa for Ouratea lucens Kunth. Engl.) to highly resistant to cavitation (P50=-4.1 MPa for Psychotria horizontalis Sw.), with shallow-rooted species being the most resistant. Among the tree species, those characteristic of waterlogged soils, Carapa guianensis Aubl., Prioria copaifera Griseb. and Ficus citrifolia Mill., were the most vulnerable to cavitation (P50=-0.8 to -1.6 MPa). The wet-season, deciduous tree, Cordia alliodora (Ruiz and Pav.) Oken., had resistant xylem (P50=-3.2 MPa), whereas the dry-season, deciduous tree, Bursera simaruba (L.) Sarg. was among the most vulnerable to cavitation (P50=-0.8 MPa) of the species studied. For eight out of the nine study species, previously reported minimum seasonal leaf water potentials measured in the field during periods of drought correlated with our P50 and P75 values. Rooting depth, deciduousness, soil type and growth habit might also contribute to desiccation tolerance. Our results support the functional dependence of drought tolerance on xylem resistance to cavitation.