Xylem Hydraulic Conductivity Research Articles

Long-term impact of Ophiostoma novo-ulmi on leaf traits and transpiration of branches in the Dutch elm hybrid 'Dodoens'.

To better understand the long-term impact of Ophiostoma novo-ulmi Brasier on leaf physiology in 'Dodoens', a Dutch elm disease-tolerant hybrid, measurements of leaf area, leaf dry mass, petiole anatomy, petiole hydraulic conductivity, leaf and branch water potential, and branch sap flow were performed 3 years following an initial artificial inoculation. Although fungal hyphae were detected in fully expanded leaves, neither anatomical nor morphological traits were affected, indicating that there was no impact from the fungal hyphae on the leaves during leaf expansion. In contrast, however, infected trees showed both a lower transpiration rate of branches and a lower sap flow density. The long-term persistence of fungal hyphae inside vessels decreased the xylem hydraulic conductivity, but stomatal regulation of transpiration appeared to be unaffected as the leaf water potential in both infected and non-infected trees was similarly driven by the transpirational demands. Regardless of the fungal infection, leaves with a higher leaf mass per area ratio tended to have a higher leaf area-specific conductivity. Smaller leaves had an increased number of conduits with smaller diameters and thicker cell walls. Such a pattern could increase tolerance towards hydraulic dysfunction. Measurements of water potential and theoretical xylem conductivity revealed that petiole anatomy could predict the maximal transpiration rate. Three years following fungal inoculation, phenotypic expressions for the majority of the examined traits revealed a constitutive nature for their possible role in Dutch elm disease tolerance of 'Dodoens' trees.

Exploiting water versus tolerating drought: water-use strategies of trees in a secondary successional tropical dry forest.

In seasonal plant communities where water availability changes dramatically both between and within seasons, understanding the mechanisms that enable plants to exploit water pulses and to survive drought periods is crucial. By measuring rates of physiological processes, we examined the trade-off between water exploitation and drought tolerance among seedlings of trees of a tropical dry forest, and identified biophysical traits most closely associated with plant water-use strategies. We also explored whether early and late secondary successional species occupy different portions of trade-off axes. As predicted, species that maintained carbon capture, hydraulic function and leaf area at higher plant water deficits during drought had low photosynthetic rates, xylem hydraulic conductivity and growth rate under non-limiting water supply. Drought tolerance was associated with more dense leaf, stem and root tissues, whereas rapid resource acquisition was associated with greater stem water storage, larger vessel diameter and larger leaf area per mass invested. We offer evidence that the water exploitation versus drought tolerance trade-off drives species differentiation in the ability of tropical dry forest trees to deal with alternating water-drought pulses. However, we detected no evidence of strong functional differentiation between early and late successional species along the proposed trade-off axes, suggesting that the environmental gradient of water availability across secondary successional habitats in the dry tropics does not filter out physiological strategies of water use among species, at least at the seedling stage.

<em>Vaccinium gaultherioides:</em> Another insight into water relations of alpine dwarf shrubs

Dwarf shrubs exhibit different requirements for a safe and efficient water supply compared to trees due their basitonic branching and low growth height. Though, only few studies dealt with the hydraulics of this growth form. Here we report key hydraulic parameters (vulnerability to drought-induced embolism, xylem hydraulic conductivity, cell osmotic potential, potential at turgor loss point) and related wood anatomical traits for Vaccinium gaultherioides, a wide-spread species in the European Alps. The results affirm the current knowledge, by indicating a relatively risky hydraulic strategy with low hydraulic safety compared to alpine trees and osmotic properties connected to the species’ soil humidity requirements.

Bark water uptake promotes localized hydraulic recovery in coastal redwood crown.

Coastal redwood (Sequoia sempervirens), the world's tallest tree species, rehydrates leaves via foliar water uptake during fog/rain events. Here we examine if bark also permits water uptake in redwood branches, exploring potential flow mechanisms and biological significance. Using isotopic labelling and microCT imaging, we observed that water entered the xylem via bark and reduced tracheid embolization. Moreover, prolonged bark wetting (16 h) partially restored xylem hydraulic conductivity in isolated branch segments and whole branches. Partial hydraulic recovery coincided with an increase in branch water potential from about -5.5 ± 0.4 to -4.2 ± 0.3 MPa, suggesting localized recovery and possibly hydraulic isolation. As bark water uptake rate correlated with xylem osmotic potential (R(2) = 0.88), we suspect a symplastic role in transferring water from bark to xylem. Using historical weather data from typical redwood habitat, we estimated that bark and leaves are wet more than 1000 h per year on average, with over 30 events being sufficiently long (>24 h) to allow for bark-assisted hydraulic recovery. The capacity to uptake biologically meaningful volumes of water via bark and leaves for localized hydraulic recovery throughout the crown during rain/fog events might be physiologically advantageous, allowing for relatively constant transpiration.

The chemical identity of intervessel pit membranes in Acer challenges hydrogel control of xylem hydraulic conductivity.

Ion-mediated enhancement of the hydraulic conductivity of xylem tissue (i.e. the ionic effect) has been reported for various angiosperm species. One explanation of the ionic effect is that it is caused by the swelling and shrinking of intervessel pit membranes due to the presence of pectins and/or other cell-wall matrix polymers such as heteroxylans or arabinogalactan-proteins (AGPs) that may contain acidic sugars. Here, we examined the ionic effect for six Acer species and their pit membrane chemistry using immunocytochemistry, including antibodies against glycoproteins. Moreover, anatomical features related to the bordered pit morphology and vessel dimensions were investigated using light and electron microscopy. The ionic effect varied from 18 % (± 9) to 32 % (± 13). Epitopes of homogalacturonan (LM18) and xylan (LM11) were not detected in intervessel pit membranes. Negative results were also obtained for glycoproteins (extensin: LM1, JIM20; AGP glycan: LM2), although AGP (JIM13)-related epitopes were detected in parenchyma cells. The mean vessel length was significantly correlated with the magnitude of the ionic effect, unlike other pit or vessel-related characteristics. Our results suggest that intervessel pit membranes of Acer are unlikely to contain pectic or other acidic polysaccharides. Therefore, alternative explanations should be tested to clarify the ionic effect.

Functional traits variation explains the distribution of Aextoxicon punctatum (Aextoxicaceae) in pronounced moisture gradients within fog-dependent forest fragments.

Climate change and fragmentation are major threats to world forests. Understanding how functional traits related to drought tolerance change across small-scale, pronounced moisture gradients in fragmented forests is important to predict species’ responses to these threats. In the case of Aextoxicon punctatum, a dominant canopy tree in fog-dependent rain forest patches in semiarid Chile, we explored how the magnitude, variability and correlation patterns of leaf and xylem vessel traits and hydraulic conductivity varied across soil moisture (SM) gradients established within and among forest patches of different size, which are associated with differences in tree establishment and mortality patterns. Leaf traits varied across soil-moisture gradients produced by fog interception. Trees growing at drier leeward edges showed higher leaf mass per area, trichome and stomatal density than trees from the wetter core and windward zones. In contrast, xylem vessel traits (vessels diameter and density) did not vary producing loss of hydraulic conductivity at drier leeward edges. We also detected higher levels of phenotypic integration and variability at leeward edges. The ability of A. punctatum to modify leaf traits in response to differences in SM availability established over short distances (<500 m) facilitates its persistence in contrasting microhabitats within forest patches. However, xylem anatomy showed limited plasticity, which increases cavitation risk at leeward edges. Greater patch fragmentation, together with fluctuations in irradiance and SM in small patches, could result in higher risk of drought-related tree mortality, with profound impacts on hydrological balances at the ecosystem scale.

Responses and adaptation of xylem hydraulic conductivity to salt stress in Populus euphratica

Responses and adaptation of xylem hydraulic conductivity to salt stress in Populus euphratica

Leaf morphology of 40 evergreen and deciduous broadleaved subtropical tree species and relationships to functional ecophysiological traits.

We explored potential of morphological and anatomical leaf traits for predicting ecophysiological key functions in subtropical trees. We asked whether the ecophysiological parameters stomatal conductance and xylem cavitation vulnerability could be predicted from microscopy leaf traits. We investigated 21 deciduous and 19 evergreen subtropical tree species, using individuals of the same age and from the same environment in the Biodiversity-Ecosystem Functioning experiment at Jiangxi (BEF-China). Information-theoretic linear model selection was used to identify the best combination of morphological and anatomical predictors for ecophysiological functions. Leaf anatomy and morphology strongly depended on leaf habit. Evergreen species tended to have thicker leaves, thicker spongy and palisade mesophyll, more palisade mesophyll layers and a thicker subepidermis. Over 50% of all evergreen species had leaves with multi-layered palisade parenchyma, while only one deciduous species (Koelreuteria bipinnata) had this. Interactions with leaf habit were also included in best multi-predictor models for stomatal conductance (gs ) and xylem cavitation vulnerability. In addition, maximum gs was positively related to log ratio of palisade to spongy mesophyll thickness. Vapour pressure deficit (vpd) for maximum gs increased with the log ratio of palisade to spongy mesophyll thickness in species having leaves with papillae. In contrast, maximum specific hydraulic conductivity and xylem pressure at which 50% loss of maximum specific xylem hydraulic conductivity occurred (Ψ50 ) were best predicted by leaf habit and density of spongy parenchyma. Evergreen species had lower Ψ50 values and lower maximum xylem hydraulic conductivities. As hydraulic leaf and wood characteristics were reflected in structural leaf traits, there is high potential for identifying further linkages between morphological and anatomical leaf traits and ecophysiological responses.

Linking xylem hydraulic conductivity and vulnerability to the leaf economics spectrum--a cross-species study of 39 evergreen and deciduous broadleaved subtropical tree species.

While the fundamental trade-off in leaf traits related to carbon capture as described by the leaf economics spectrum is well-established among plant species, the relationship of the leaf economics spectrum to stem hydraulics is much less known. Since carbon capture and transpiration are coupled, a close connection between leaf traits and stem hydraulics should be expected. We thus asked whether xylem traits that describe drought tolerance and vulnerability to cavitation are linked to particular leaf traits. We assessed xylem vulnerability, using the pressure sleeve technique, and anatomical xylem characteristics in 39 subtropical tree species grown under common garden conditions in the BEF-China experiment and tested for correlations with traits related to the leaf economics spectrum as well as to stomatal control, including maximum stomatal conductance, vapor pressure deficit at maximum stomatal conductance and vapor pressure deficit at which stomatal conductance is down-regulated. Our results revealed that specific xylem hydraulic conductivity and cavitation resistance were closely linked to traits represented in the leaf economic spectrum, in particular to leaf nitrogen concentration, as well as to log leaf area and leaf carbon to nitrogen ratio but not to any parameter of stomatal conductance. The study highlights the potential use of well-known leaf traits from the leaf economics spectrum to predict plant species' drought resistance.

How do drought and warming influence survival and wood traits of Picea mariana saplings?

Warming and drought will occur with increased frequency and intensity at high latitudes in the future. How heat and water stress can influence tree mortality is incompletely understood. The aim of this study was to evaluate how carbon resources, stem hydraulics, and wood anatomy and density determine the ability of black spruce saplings to survive daytime or night-time warming (+ 6 °C in comparison with control) in combination with a drought period. Plant water relations, the dynamics of non-structural carbohydrates and starch, mortality rate, and wood anatomy and density of saplings were monitored. Warming, in conjunction with 25 d of water deficit, increased sapling mortality (10% and 20% in night-time and daytime warming, respectively) compared with the control conditions (0.8%). Drought substantially decreased gas exchange, and also pre-dawn and mid-day leaf water potential to values close to -3MPa which probably induced xylem embolism (xylem air entry point, P₁₂, being on average around -3MPa for this species). In addition, the recovery of gas exchange never reached the initial pre-stress levels, suggesting a possible loss of xylem hydraulic conductivity associated with cavitation. Consequently, mortality may be due to xylem hydraulic failure. Warmer temperatures limited the replenishment of starch reserves after their seasonal minimum. Lighter wood was formed during the drought period, reflecting a lower carbon allocation to cell wall formation, preventing the adaptation of the hydraulic system to drought. Saplings of black spruce experienced difficulty in adapting under climate change conditions, which might compromise their survival in the future.

Are leaves 'freewheelin'? Testing for a wheeler-type effect in leaf xylem hydraulic decline.

A recent study found that cutting shoots under water while xylem was under tension (which has been the standard protocol for the past few decades) could produce artefactual embolisms inside the xylem, overestimating hydraulic vulnerability relative to shoots cut under water after relaxing xylem tension (Wheeler et al. 2013). That study also raised the possibility that such a 'Wheeler effect' might occur in studies of leaf hydraulic vulnerability. We tested for such an effect for four species by applying a modified vacuum pump method to leaves with minor veins severed, to construct leaf xylem hydraulic vulnerability curves. We tested for an impact on leaf xylem hydraulic conductance (Kx ) of cutting the petiole and minor veins under water for dehydrated leaves with xylem under tension compared with dehydrated leaves after previously relaxing xylem tension. Our results showed no significant 'cutting artefact' for leaf xylem. The lack of an effect for leaves could not be explained by narrower or shorter xylem conduits, and may be due to lesser mechanical stress imposed when cutting leaf petioles, and/or to rapid refilling of emboli in petioles. These findings provide the first validation of previous measurements of leaf hydraulic vulnerability against this potential artefact.

Sap flow-based quantitative indication of progression of Dutch elm disease after inoculation with Ophiostoma novo-ulmi

The rate of progression of Dutch elm disease can be continuously and quantitatively estimated from sap flow measurements. Response of sap flow to inoculation with Ophiostoma novo-ulmi, a causal agent which causes vascular mycosis called Dutch elm disease, was studied in a field experiment comprised of 4-year-old wych elm trees (Ulmus glabra). Sap flow was measured on inoculated trees using the trunk heat balance method with external heating (EMS 62, Czech Republic) throughout the experiment. The first detectable symptoms of reduction in sap flow occurred 6 days after inoculation and all inoculated trees died within 16 days. Our experiment confirmed the ability of O. novo-ulmi to quickly kill young elm trees. The disease progressed faster than in previous experiments utilizing O. ulmi. To the best of our knowledge, this is the first experiment using sap flow measurements on trees inoculated by O. novo-ulmi. The trunk heat balance sap flow method is an effective non-invasive tool for continuous quantitative monitoring of the progression of vascular tree diseases, and show increased potential for field and greenhouse studies on changes in xylem hydraulic conductivity in a wide range of broadleaved and coniferous tree species.

Leaf and stem physiological responses to summer and winter extremes of woody species across temperate ecosystems

Winter cold limits temperate plant performance, as does summer water stress in drought‐prone ecosystems. The relative impact of seasonal extremes on plant performance has received considerable attention for individual systems. An integrated study compiling the existing literature was needed to identify overall trends. First, we conducted a meta‐analysis of the impacts of summer and winter on ecophysiology for three woody plant functional types (winter deciduous angiosperms, evergreen angiosperms and conifers), including data for 210 records from 75 studies of ecosystems with and without summer drought across the temperate zone. Second, we tested predictions by conducting a case study in a drought‐prone Mediterranean ecosystem subject to winter freezing. As indicators of physiological response of leaves and xylem to seasonal stress, we focused on stomatal conductance (gs), percent loss of stem xylem hydraulic conductivity (PLC) and photochemical efficiency of photosystem II (Fv/Fm). Our meta‐analysis showed that in ecosystems without summer drought, gs was higher during summer than winter. By contrast, in drought‐prone ecosystems many species maintained open stomata during winter, with potential strong consequences for plant carbon gain over the year. Further, PLC tended to increase and Fv/Fm to decrease from summer to winter for most functional types and ecosystems due to low temperatures. Overall, deciduous angiosperms were most sensitive to climatic stress. Leaf gas exchange and stem xylem hydraulics showed a coordinated seasonal response at ecosystems without summer drought. In our Mediterranean site subjected to winter freezing the species showed similar responses to those typically found for ecosystems without summer drought. We conclude that winter stress is most extreme for systems without summer drought and systems with summer drought and winter freezing, and less extreme for drought‐prone systems without freezing. In all cases the evergreen species show less pronounced seasonal responses in both leaves and stems than deciduous species.

Soil water deficit decreases xylem conductance efficiency relative to leaf area and mass in the apple

It is generally postulated that at the tree scale a drought-related decrease in hydraulic conductance is balanced by a decrease of leaf area. We hypothesized that, at the individual leaf scale, drought affects the allometry between leaf area or mass and hydraulics, leading to a non-linear relationships between these traits. The study was conducted on well-watered and on water-stressed shoots of several apple genotypes covering an extended range of leaf area. Working on dried leaves, we measured leaf lamina area and mass and analyzed their relationships with the maximal xylem hydraulic conductance of the water pathway through the parent shoot and the petiole connected to the leaf lamina. Drought decreased leaf area and mass in absolute values. It also changes the allometric relationships between these two variables: for a same decrease of leaf dry mass the water-stressed shoot had a lower decrease of leaf dry area than the well-watered shoot. Our study also showed that drought affected the stem-to-petiole hydraulics with a higher hydraulic efficiency in the well-watered shoot compared to the water-stressed shoot. We discuss that, compared to the well-watered condition, drought not only decreased leaf size, but also reduced xylem efficiency through the stem-to-petiole pathway with regard to the leaf area and mass supplied.

Recent advances in tree hydraulics highlight the ecological significance of the hydraulic safety margin

Drought-induced cavitation resistance varies considerably between tree species and forest ecosystems (Maherali et al., 2004; Delzon et al., 2010) and is closely linked to survival under severe drought in both conifers (Brodribb & Cochard, 2009; Brodribb et al., 2010) and angiosperms (Kursar et al., 2009; Anderegg et al., 2012; Barigah et al., 2013; Urli et al., 2013). Choat et al. (2012) recently reported that most trees operate very close to their threshold of cavitation, leaving them potentially vulnerable to drought-induced mortality in a warmer/drier world (Engelbrecht, 2012). Indeed, species growing in dry environments are more resistant to droughtinduced cavitation (more negative water potential at 50% cavitation, P50) but experience a more negative minimumwater potential (Pmin) than those growing in wet environments. The so-called hydraulic safety margin, the difference between the level of water stress experienced by a species in the field (Pmin) and the level of water stress leading to hydraulic failure, is, therefore, remarkably narrow, whatever the forest species and biome considered (Choat et al., 2012). This pattern provides clues to the global droughtinduced mortality currently observed, even in very wet environments, such as tropical forests (Allen et al., 2010). Klein et al. (2014) play down the functional significance of the hydraulic safety margin in the vulnerability of forests to drought, pointing out the important role played by additional mechanisms, such as the ability of trees to repair embolism. While it is obvious that drought-induced forest dieback is a complex process involving a number of biotic and abiotic factors, we would like to draw the attention of scientists to the state of evidence for embolism repair, thereby guiding research on tree drought resistance into the most relevant and fruitful directions

Coping with drought-induced xylem cavitation: coordination of embolism repair and ionic effects in three Mediterranean evergreens

Embolism repair and ionic effects on xylem hydraulic conductance have been documented in different tree species. However, the diurnal and seasonal patterns of both phenomena and their actual role in plants' responses to drought-induced xylem cavitation have not been thoroughly investigated. This study provides experimental evidence of the ability of three Mediterranean species to maintain hydraulic function under drought stress by coordinating the refilling of xylem conduits and ion-mediated enhancement of stem hydraulic conductance (K stem). Vessel grouping indices and starch content in vessel-associated parenchyma cells were quantified to verify eventual correlations with ionic effects and refilling, respectively. Experiments were performed on stems of Ceratonia siliqua L., Olea europaea L. and Laurus nobilis L. Seasonal, ion-mediated changes in K stem (ΔK stem) and diurnal and/or seasonal embolism repair were recorded for all three species, although with different temporal patterns. Field measurements of leaf specific stem hydraulic conductivity showed that it remained quite constant during the year, despite changes in the levels of embolism. Starch content in vessel-associated parenchyma cells changed on diurnal and seasonal scales in L. nobilis and O. europaea but not in C. siliqua. Values of ΔK stem were significantly correlated with vessel multiple fraction values (the ratio of grouped vessels to total number of vessels). Our data suggest that the regulation of xylem water transport in Mediterranean plants relies on a close integration between xylem refilling and ionic effects. These functional traits apparently play important roles in plants' responses to drought-induced xylem cavitation.

Does short-term potassium fertilization improve recovery from drought stress in laurel?

Xylem hydraulic conductance varies in response to changes in sap solute content, and in particular of potassium (K(+)) ion concentration. This phenomenon, known as the 'ionic effect', is enhanced in embolized stems, where it can compensate for cavitation-induced loss of hydraulic conductance. Previous studies have shown that in well-watered laurel plants (Laurus nobilis L.), potassium concentration of the xylem sap and plant hydraulic conductance increased 24 h after fertilization with KCl. The aim of this work was to test whether water-stressed laurel plants, grown under low potassium availability, could recover earlier from stress when irrigated with a KCl solution instead of potassium-free water. Two-year-old potted laurel seedlings were subjected to water stress by suspending irrigation until leaf conductance to water vapour (g(L)) dropped to ∼30% of its initial value and leaf water potential (ψ(L)) reached the turgor loss point (ψ(TLP)). Plants were then irrigated either with water or with 25 mM KCl and monitored for water status, gas exchange and plant hydraulics recovery at 3, 6 and 24 h after irrigation. No significant differences were found between the two experimental groups in terms of ψ(L), g(L), plant transpiration, plant hydraulic conductance or leaf-specific shoot hydraulic conductivity. Analysis of xylem sap potassium concentration showed that there were no significant differences between treatments, and potassium levels were similar to those of potassium-starved but well-watered plants. In conclusion, potassium uptake from the soil solution and/or potassium release to the xylem appeared to be impaired in water-stressed plants, at least up to 24 h after relief from water stress, so that fertilization after the onset of stress did not result in any short-term advantage for recovery from drought.

“Plastic” and “static” behavior of vessel-anatomical features in Oriental beech (Fagus orientalis Lipsky) in view of xylem hydraulic conductivity

Key message Xylem anatomical traits can be categorized into two groups: plastic properties which show a high inter-annual variability, and static characteristics which vary in a more conservative range.

GEL OCCLUSION IN THE XYLEM VESSELS OF CUT ACACIA HOLOSERICEA FOLIAGE STEMS

The short commercial vase life of cut Acacia holosericea foliage is potentially the result of physiologically-mediated xylem vessel occlusion. Xylem-occluding gels are considered to be a defence and wound healing mechanism. In ∼45 cm-long cut A. holosericea foliage stems, occluding gel secretions were observed in xylem vessel lumens at the cut end and for up to 35 cm distally from the cut end. The gels were deposited soon after the stems were detached (mechanically wounded) and they apparently matured over time. Early gels were transparent and colourless in reaction with phloroglucinol-HCl (P-HCl). The staining of late gels with P-HCl was typically pink to cherry red, amber and yellow, or light brown to dark brown. The deposition of gels evidently reduced the xylem hydraulic conductance (Kh) of cut A. holosericea foliage stems.

Species Variation in Water Relations and Xylem Vulnerability to Cavitation at a Forest-Woodland Ecotone

Xylem vulnerability to cavitation and response of water potential (), stomatal conductance (g s), and net photosynthesis (P n) to drought are potentially important mechanisms of drought resistance. We compared , g s, Pn, and cavitation vulnerability of shoot and root xylem among co-occurring ponderosa pine (Pinus ponderosa var. scopulorum Dougl. Ex Laws.), pinyon pine (Pinus edulis Engelm.), and Utah juniper (Juniperus osteosperma (Torr.) Little) at a forest-woodland ecotonal site in northern Arizona to elucidate drought resistance mechanisms of these species. Juniper shoots partly regulated during drought via stomatal closure, but regulation was weaker than that for ponderosa and pinyon pines, which had similar water relations and P n responses to drought. Midday g s and Pn during summer drought were positive for juniper (gs 14.3 mmol m 2 s 1 , Pn 1.23 mol m 2 s 1 ) but near zero for ponderosa (gs 0.7 mmol m 2 s 1 , Pn 0.02 mol m 2 s 1 ) and pinyon (gs 1.5 mmol m 2 s 1 , Pn 0.18 mol m 2 s 1 ) pines. Cavitation vulnerability of shoots and roots was lower for juniper than for both pines. The water potential inducing 50% loss in xylem hydraulic conductivity ( 50) for juniper was 5.0 MPa more negative for shoots and 3.9 MPa more negative for roots compared with the respective tissues of the pine species. Pinyon pine ( 50 2.71 MPa) was slightly more vulnerable to cavitation than ponderosa pine ( 50 3.42 MPa) for shoots, whereas root vulnerability was similar for both pines ( 50 1.69 MPa for pinyon; 1.98 MPa for ponderosa). Roots of all species were more vulnerable to cavitation than shoots. Our results show an important role of cavitation vulnerability in the greater drought resistance of Utah juniper than pinyon and ponderosa pines but not for the presumed greater drought resistance of pinyon pine than ponderosa pine. FOR .S CI. 59(5):524-535.