Functional xylem anatomy of aspen exhibits greater change due to insect defoliation than to drought.

The study of tree rings can reveal long-term records of a tree's response to the environment. This dendroecological approach, when supplemented with finer-scale observations of the xylem anatomy, can provide novel information about a tree's year-to-year anatomical and hydraulic adjustments. Here we use this method in aspen (Populus tremuloides Michx.) to identify xylem response to drought and insect defoliation. Surprisingly, we found that precipitation influenced vessel diameter mostly in the trees' youth, while this correlation was less pronounced at maturity. This is likely due to a reduction in stress the stand experiences as it ages, and reflects an ability to mediate drought stress as trees mature. Defoliation events caused consistent and profound changes in fiber anatomy likely leading to reduced structural support to vessels. We therefore expect that in years of defoliation trees may be vulnerable to drought-induced cavitation when leaf area recovers. This study highlights how the inclusion of cellular level measurements in tree ring studies provides additional information on how stress events may alter tree functioning through alterations in structure.

Xylem function of arid-land shrubs from California, USA: an ecological and evolutionary analysis.

Xylem traits were examined among 22 arid-land shrub species, including measures of vessel dimensions and pit area. These structural measures were compared with the xylem functional traits of transport efficiency and safety from cavitation. The influence of evolution on trait relationships was examined using phylogenetic independent contrasts (PICs). A trade-off between xylem safety and efficiency was supported by a negative correlation between vessel dimensions and cavitation resistance. Pit area was correlated with cavitation resistance when cross species data were examined, but PICs suggest that these traits have evolved independently of one another. Differences in cavitation resistance that are not explained by pit area may be related to differences in pit membrane properties or the prevalence of tracheids, the latter of which may alter pit area through the addition of vessel-to-tracheid pits or through changes in xylem conduit connectivity. Some trait relationships were robust regardless of species ecology or evolutionary history. These trait relationships are likely to be the most valuable in predictive models that seek to examine anatomical and functional trait relationships among extant and fossil woods and include the relationship among hydraulic conductivity and vessel diameter, between vessel diameter and vessel length, and between hydraulic conductivity and wood density.

Adjustments in hydraulic architecture of Pinus palustris maintain similar stomatal conductance in xeric and mesic habitats.

We investigated relationships between whole-tree hydraulic architecture and stomatal conductance in Pinus palustris Mill. (longleaf pine) across habitats that differed in soil properties and habitat structure. Trees occupying a xeric habitat (characterized by sandy, well-drained soils, higher nitrogen availability and lower overstory tree density) were shorter in stature and had lower sapwood-to-leaf area ratio (A(S):A(L)) than trees in a mesic habitat. The soil-leaf water potential gradient (psiS - psiL) and leaf-specific hydraulic conductance (kL) were similar between sites, as was tissue-specific hydraulic conductivity (Ks) of roots. Leaf and canopy stomatal conductance (gs and Gs, respectively) were also similar between sites, and they tended to be somewhat higher at the xeric site during morning hours when vapour pressure deficit (D) was low. A hydraulic model incorporating tree height, A(S):A(L) and psiS-psiL accurately described the observed variation in individual tree G(Sref) (G(S) at D = 1 kPa) across sites and indicated that tree height was an important determinant of G(Sref) across sites. This, combined with a 42% higher root-to-leaf area ratio (A(R):A(L)) at the xeric site, suggests that xeric site trees are hydraulically well equipped to realize equal--and sometimes higher potential for conductance compared with trees on mesic sites. However, a slightly more sensitive stomatal closure response to increasing D observed in xeric site trees suggests that this potential for higher conductance may only be reached when D is low and when the capacity of the hydraulic system to supply water to foliage is not greatly challenged.

Water deficits and hydraulic limits to leaf water supply.

Many aspects of plant water use -- particularly in response to soil drought -- may have as their basis the alteration of hydraulic conductance from soil to canopy. The regulation of plant water potential (Psi) by stomatal control and leaf area adjustment may be necessary to maximize water uptake on the one hand, while avoiding loss of hydraulic contact with the soil water on the other. Modelling the changes in hydraulic conductance with pressure gradients in the continuum allows the prediction of water use as a function of soil environment and plant architectural and xylem traits. Large differences in water use between species can be attributed in part to differences in their 'hydraulic equipment' that is presumably optimized for drawing water from a particular temporal and spatial niche in the soil environment. A number of studies have identified hydraulic limits as the cause of partial or complete foliar dieback in response to drought. The interactions between root:shoot ratio, rooting depth, xylem properties, and soil properties in influencing the limits to canopy water supply can be used to predict which combinations should optimize water use in a given circumstance. The hydraulic approach can improve our understanding of the coupling of canopy processes to soil environment, and the adaptive significance of stomatal behaviour.

Cavitation fatigue. Embolism and refilling cycles can weaken the cavitation resistance of xylem.

Although cavitation and refilling cycles could be common in plants, it is unknown whether these cycles weaken the cavitation resistance of xylem. Stem or petiole segments were tested for cavitation resistance before and after a controlled cavitation-refilling cycle. Cavitation was induced by centrifugation, air drying of shoots, or soil drought. Except for droughted plants, material was not significantly water stressed prior to collection. Cavitation resistance was determined from "vulnerability curves" showing the percentage loss of conductivity versus xylem pressure. Two responses were observed. "Resilient" xylem (Acer negundo and Alnus incana stems) showed no change in cavitation resistance after a cavitation-refilling cycle. In contrast, "weakened" xylem (Populus angustifolia, P. tremuloides, Helianthus annuus stems, and Aesculus hippocastanum petioles) showed considerable reduction in cavitation resistance. Weakening was observed whether cavitation was induced by centrifugation, air dehydration, or soil drought. Observations from H. annuus showed that weakening was proportional to the embolism induced by stress. Air injection experiments indicated that the weakened response was a result of an increase in the leakiness of the vascular system to air seeding. The increased air permeability in weakened xylem could result from rupture or loosening of the cellulosic mesh of interconduit pit membranes during the water stress and cavitation treatment.

Influence of soil porosity on water use in Pinus taeda.

We analyzed the hydraulic constraints imposed on water uptake from soils of different porosities in loblolly pine (Pinus taeda L.) by comparing genetically related and even-aged plantations growing in loam versus sand soil. Water use was evaluated relative to the maximum transpiration rate (E crit) allowed by the soil-leaf continuum. We expected that trees on both soils would approach E crit during drought. Trees in sand, however, should face greater drought limitation because of steeply declining hydraulic conductivity in sand at high soil water potential (Ψ S). Transport considerations suggest that trees in sand should have higher root to leaf area ratios (A R:A L), less negative leaf xylem pressure (Ψ L), and be more vulnerable to xylem cavitation than trees in loam. The A R:A L was greater in sand versus loam (9.8 vs 1.7, respectively). This adjustment maintained about 86% of the water extraction potential for both soils. Trees in sand were more deeply rooted (>1.9 m) than in loam (95% of roots <0.2 m), allowing them to shift water uptake to deeper layers during drought and avoid hydraulic failure. Midday Ψ L was constant for days of high evaporative demand, but was less negative in sand (-1.6 MPa) versus loam (-2.1 MPa). Xylem was more vulnerable to cavitation in sand versus loam trees. Roots in both soils were more vulnerable than stems, and experienced the greatest predicted loss of conductivity during drought. Trees on both soils approached E crit during drought, but at much higher Ψ S in sand (<-0.4 MPa) than in loam (<-1.0 MPa). Results suggest considerable phenotypic plasticity in water use traits for P. taeda which are adaptive to differences in soil porosity.

The relationship between xylem conduit diameter and cavitation caused by freezing.

The centrifuge method for measuring the resistance of xylem to cavitation by water stress was modified to also account for any additional cavitation that might occur from a freeze-thaw cycle. A strong correlation was found between cavitation by freezing and mean conduit diameter. On the one extreme, a tracheid-bearing conifer and diffuse-porous angiosperms with small-diameter vessels (mean diameter <30 µm) showed no freezing-induced cavitation under modest water stress (xylem pressure = -0.5 MPa), whereas species with larger diameter vessels (mean >40 µm) were nearly completely cavitated under the same conditions. Species with intermediate mean diameters (30-40 µm) showed partial cavitation by freezing. These results are consistent with a critical diameter of 44 µm at or above which cavitation would occur by a freeze-thaw cycle at -0.5 MPa. As expected, vulnerability to cavitation by freezing was correlated with the hydraulic conductivity per stem transverse area. The results confirm and extend previous reports that small-diameter conduits are relatively resistant to cavitation by freezing. It appears that the centrifuge method, modified to include freeze-thaw cycles, may be useful in separating the interactive effects of xylem pressure and freezing on cavitation.