Defoliation constrains xylem and phloem functionality.

Insect defoliation contributes to tree mortality under drought conditions. Defoliation-induced alterations to the vascular transport structure may increase tree vulnerability to drought; however, this has been rarely studied. To evaluate the response of tree vascular function following defoliation, 2-year-old balsam poplar were manually defoliated, and both physiological and anatomical measurements were made after allowing for re-foliation. Hydraulic conductivity measurements showed that defoliated trees had both increased vulnerability to embolism and decreased water transport efficiency, likely due to misshapen xylem vessels. Anatomical measurements revealed novel insights into defoliation-induced alterations to the phloem. Phloem sieve tube diameter was reduced in the stems of defoliated trees, suggesting reduced transport capability. In addition, phloem fibers were absent, or reduced in number, in stems, shoot tips and petioles of new leaves, potentially reducing the stability of the vascular tissue. Results from this study suggest that the defoliation leads to trees with increased risk for vascular dysfunction and drought-induced mortality through alterations in the vascular structure, and highlights a route through which carbon limitation can influence hydraulic dysfunction.

Re-establishment of hummock topography promotes tree regeneration on highly disturbed moderate-rich fens.

Winter exploration of oil sands deposits underlying wooded fens mostly eliminates the hummock-hollow topography on drilling pads and the ice roads leading to them, after their abandonment in spring. Recovery of black spruce (Picea mariana (P. Mill.) B.S.P.) and tamarack (Larix laricina (Du Roi) K. Koch) on these disturbed peatlands is thought to depend on the recovery of hummock topography. In late winter, numerous large blocks of frozen peat (1.5 × 1.5 m) were lifted out of the flattened drilling pads and positioned beside their excavated hollows; this was done on six temporary pads. Four years later, the condition of the mounds and the regeneration of conifers from natural seed dispersal were assessed on these elevated mounds compared to adjacent flattened areas of the pads. Then, conifer seedling density was more than five times higher on elevated spots than the mostly flat, flood-prone areas between them, and seedling density was positively related to mound height and strength of seed source. Higher mounds tended to have larger seedlings. Mounds on some of the pads were heavily eroded down; these pads had peat with higher humification, and operationally these pads were also treated in late winter when peat was thawing and fractured into pieces during mound construction. Developing a large volume of elevated substrate that persists until natural hummock-forming mosses can establish is thought necessary for tree recruitment and the recovery of the habitat for the threatened woodland caribou of this region.

Drought-induced xylem pit membrane damage in aspen and balsam poplar.

Drought induces an increase in a tree's vulnerability to a loss of its hydraulic conductivity in many tree species, including two common in western Canada, trembling aspen (Populus tremuloides) and balsam poplar (Populus balsamifera). Termed 'cavitation fatigue' or 'air-seeding fatigue', the mechanism of this phenomenon is not well understood, but hypothesized to be a result of damage to xylem pit membranes. To examine the validity of this hypothesis, the effect of drought on the porosity of pit membranes in aspen and balsam poplar was investigated. Controlled drought and bench dehydration treatments were used to induce fatigue and scanning electron microscopy (SEM) was used to image pit membranes for relative porosity evaluations from air-dried samples after ethanol dehydration. A significant increase in the diameter of the largest pore was found in the drought and dehydration treatments of aspen, while an increase in the percentage of porous pit membranes was found in the dehydration treatments of both species. Additionally, the location of the largest pore per pit membrane was observed to tend toward the periphery of the membrane.

Daytime and nighttime wind differentially affects hydraulic properties and thigmomorphogenic response of poplar saplings.

This study tested how wind in daytime and nighttime affects hydraulic properties and thigmomorphogenic response of poplar saplings. It shows that wind in daytime interrupted water balance of poplar plants by aggravating cavitation in the stem xylem under high xylem tension in the daytime, reducing water potential in midday and hence reducing gas exchange, including stomatal conductance and CO2 assimilation. The wind blowing in daytime significantly reduced plant growth, including height, diameter, leaf size, leaf area, root and whole biomass, whereas wind blowing in nighttime only caused a reduction in radial and height growth at the early stage compared with the control but decreased height:diameter ratios. In summary, the interaction between wind loading and xylem tension exerted a negative impact on water balance, gas exchanges and growth of poplar plants, and wind in nighttime caused only a small thigmomorphogenic response.

Uniform versus asymmetric shading mediates crown recession in conifers.

In this study we explore the impact of asymmetrical vs. uniform crown shading on the mortality and growth of upper and lower branches within tree crowns, for two conifer species: shade intolerant lodgepole pine (Pinus contorta) and shade tolerant white spruce (Picea glauca). We also explore xylem hydraulics, foliar nutrition, and carbohydrate status as drivers for growth and expansion of the lower and upper branches in various types of shading. This study was conducted over a two-year period across 10 regenerating forest sites dominated by lodgepole pine and white spruce, in the lower foothills of Alberta, Canada. Trees were assigned to one of four shading treatments: (1), complete uniform shading of the entire tree, (2) light asymmetric shading where the lower 1/4-1/3 of the tree crown was shaded, (3) heavy asymmetric shading as in (2) except with greater light reduction and (4) control in which no artificial shading occurred and most of the entire crown was exposed to full light. Asymmetrical shading of only the lower crown had a larger negative impact on the bud expansion and growth than did uniform shading, and the effect was stronger in pine relative to spruce. In addition, lower branches in pine also had lower carbon reserves, and reduced xylem-area specific conductivity compared to spruce. For both species, but particularly the pine, the needles of lower branches tended to store less C than upper branches in the asymmetric shade, which could suggest a movement of reserves away from the lower branches. The implications of these findings correspond with the inherent shade tolerance and self-pruning behavior of these conifers and supports a carbon based mechanism for branch mortality--mediated by an asymmetry in light exposure of the crown.

Variation in carbon availability, defense chemistry and susceptibility to fungal invasion along the stems of mature trees.

If carbon (C) sinks withdraw carbohydrates as they are transported along tree stems, carbohydrate availability may depend on local sink strength and distance from sources. Defenses, including monoterpenes--a major component of resin--limit the invasibility of pines. Since carbohydrate reserves fund monoterpene synthesis, we hypothesized that monoterpene concentrations in pine stems would decrease from the crown to the lower stem, and susceptibility to fungal infection would increase. Here, we measured carbohydrate and monoterpene concentrations along the stems of lodgepole pine trees (Pinus contorta var. latifolia) before inoculating with a blue-stain fungus at different heights. After 6 wk, we assessed tree responses to fungal infection based on lesion length and carbohydrate mobilization. Concentrations of carbohydrates and monoterpenes in the phloem before inoculation decreased with distance from the crown, whereas lesion lengths after inoculation increased. However, trees mobilized sugars in response to fungal infection such that carbohydrate reserves near lesions were similar at all heights. Despite C mobilization, the lower stem was more vulnerable than the upper stem. Consistent with predictions based on sink-source relationships, vulnerability occurred where carbohydrates were less available, and likely resulted from C withdrawal by sinks higher in the supply chain.

The impact of phloem nutrients on overwintering mountain pine beetles and their fungal symbionts.

In the low nutrient environment of conifer bark, subcortical beetles often carry symbiotic fungi that concentrate nutrients in host tissues. Although bark beetles are known to benefit from these symbioses, whether this is because they survive better in nutrient-rich phloem is unknown. After manipulating phloem nutrition by fertilizing lodgepole pine trees (Pinus contorta Douglas var. latifolia), we found bolts from fertilized trees to contain more living individuals, and especially more pupae and teneral adults than bolts from unfertilized trees at our southern site. At our northern site, we found that a larger proportion of mountain pine beetle (Dendroctonus ponderosae Hopkins) larvae built pupal chambers in bolts from fertilized trees than in bolts from unfertilized trees. The symbiotic fungi of the mountain pine beetle also responded to fertilization. Two mutualistic fungi of bark beetles, Grosmannia clavigera (Rob.-Jeffr. & R. W. Davidson) Zipfel, Z. W. de Beer, & M. J. Wingf. and Leptographium longiclavatum Lee, S., J. J. Kim, & C. Breuil, doubled the nitrogen concentrations near the point of infection in the phloem of fertilized trees. These fungi were less capable of concentrating nitrogen in unfertilized trees. Thus, the fungal symbionts of mountain pine beetle enhance phloem nutrition and likely mediate the beneficial effects of fertilization on the survival and development of mountain pine beetle larvae.

Viability of forest floor and canopy seed banks in Pinus contorta var. latifolia (Pinaceae) forests after a mountain pine beetle outbreak.

PREMISE OF THE STUDY: Seed banks are important for the natural regeneration of many forest species. Most of the seed bank of serotinous lodgepole pine is found in the canopy, but after an outbreak of mountain pine beetle (MPB), a considerable forest-floor seed bank develops through the falling of canopy cones. After large-scale mortality of pine stands from MPB, however, the viability of seeds in both the canopy and the forest-floor cone bank is uncertain. METHODS: We sampled cones in five stands 3 yr after MPB (3y-MPB); five stands 6 yr after MPB (6y-MPB); and 10 stands 9 yr after MPB (9y-MPB), in central British Columbia, Canada. Seeds were extracted and viability tested using germination techniques. KEY RESULTS: Forest-floor cones had seed with high germination capacity (GC): 82% for embedded (partly buried) closed cones vs. 45% for buried partly open cones. For canopy cones, GC steeply declined about 15 yr after cone maturation and by 25 yr, GC was 50%, compared with 98% in the first year. In the 3y- and 6y-MPB stands, seeds from cones that were 7 to 9 yr old had similar GC on dead and living trees; however, seeds from the dead trees had lower vigor than seeds from living trees. CONCLUSIONS: We demonstrate for the first time that a serotinous pine can form a viable soil seed bank by cone burial, which may facilitate natural regeneration if a secondary disturbance occurs. Seeds contained in 15-yr-old cones showed a steep decline in viability, which could limit regeneration if there is a long delay before a secondary disturbance.

Seed release in serotinous lodgepole pine forests after mountain pine beetle outbreak.

There are concerns that large-scale stand mortality due to mountain pine beetle (MPB) could greatly reduce natural regeneration of serotinous Rocky Mountain (RM) lodgepole pine (Pinus contorta var. latifolia) because the closed cones are held in place without the fire cue for cone opening. We selected 20 stands (five stands each of live [control], 3 years since MPB [3-yr-MPB], 6 years since MPB [6-yr-MPB], and 9 years since MPB [9-yr-MPB] mortality) in north central British Columbia, Canada. The goal was to determine partial loss of serotiny due to fall of crown-stored cones via breakage of branches and in situ opening of canopy cones throughout the 2008 and 2009 growing seasons. We also quantified seed release by the opening of forest-floor cones, loss of seed from rodent predation, and cone burial. Trees killed by MPB three years earlier dropped approximately 3.5 times more cones via branch breakage compared to live stands. After six years, MPB-killed stands had released 45% of their canopy seed bank through cone opening, cone fall due to breakage, and squirrel predation. Further losses of canopy seed banks are expected with time since we found 9-yr-MPB stands had 38% more open canopy cones. This was countered by the development of a modest forest-floor seed bank (6% of the original canopy seed bank) from burial of cones; this seed bank may be ecologically important if a fire or anthropogenic disturbance reexposes these cones. If adequate levels of regeneration are to occur, disturbances to create seedbeds must occur shortly after tree mortality, before the seed banks are lost. Our findings also suggest that the sustained seed rain (over at least nine years) after MPB outbreak may be beneficial for population growth of ground-foraging vertebrates. Our study adds insight to the seed ecology of serotinous pines under a potentially continental-wide insect outbreak, threatening vast forests adapted to regeneration after fire. Key words: biotic disturbance; cone burial; cone opening; Dendroctonus ponderosae; ground-foraging vertebrates; mountain pine beetle; natural regeneration; Pinus contorta var. latifolia; Rocky Mountain lodgepole pine; seed banks; serotiny (canopy seed storage); Tamiasciurus hudsonicus.

Patterns of inter-annual variation in the size asymmetry of growth in Pinus banksiana.

A large body of literature suggests that asymmetric competition, where large individuals suppress the growth of smaller individuals by intercepting a disproportionate share of incoming light, is a dominant process in tree population development. This has not been examined extensively for long-lived tree species that accumulate growth over many years under varying growing conditions. Using dendrochronological techniques, we reconstructed annual growth and mortality rates at ten stands of jack pine (Pinus banksiana Lamb.) in Western Canada. We used these data to calculate an annual index of the size asymmetry of growth for each stand for the last 50 years. Jack pine is a shade-intolerant species found in even-aged monoculture stands, so the simple hypothesis is that large trees should consistently perform relatively better than small trees. Inter-annual variation in the index of size-asymmetric growth was positively associated with interannual variation in stand productivity at eight of ten sites. The size asymmetry of growth also showed a positive trend with age at eight of ten sites, even though all sites were in a period of declining leaf area. This should have reduced the intensity of asymmetric competition for light and reduced the size asymmetry of growth over time. Alternate hypotheses for this trend are (1) that physical collisions between crowns result in asymmetric competition for growing space because they are more damaging to small trees, or (2) that a differential build up of diseases in susceptible trees suppresses their growth, even in the absence of competition.

Growth-climate relationships vary with height along the stem in lodgepole pine.

This study tests the hypothesis that ring growth in the upper stem portion of trees is affected by climatic conditions differently than rings formed at breast height (1.3 m). A total of 389 trees from a network of 65 lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) sites in Alberta were examined using detailed stem analysis in order to examine interannual patterns of basal area increment and volume increment at different positions along the stem. Growth at lower sections of the bole was mainly driven by temperature and moisture conditions in the seasons prior to the growing season in the year of ring formation, while upper stem growth was more related to conditions during the year of growth, i.e., temperature in the early summer, or moisture in late winter to early spring. This translates into increased allocation of wood to the lower stem when prior late summer conditions are cool and wet, prior winters are mild (warm with little snow) and early summer conditions in the year of ring formation are hot and dry.

Hydraulic acclimation to shading in boreal conifers of varying shade tolerance.

The purpose of this study was to determine how shading affects the hydraulic and wood-anatomical characteristics of four boreal conifers (Pinus banksiana, Pinus contorta, Picea glauca and Picea mariana) that differ in shade tolerance. Plants were grown in an open field and under a deciduous-dominated overstory for 6 years. Sapwood- and leaf-area specific conductivity, vulnerability curves, and anatomical measurements (light and scanning electron microscopy) were made on leading shoots from six to nine trees of each treatment combination. There was no difference in sapwood-area specific conductivity between open-grown and understory conifers, although two of four species had larger tracheid diameters in the open. Shaded conifers appeared to compensate for small diameter tracheids by changes in pit membrane structure. Scanning electron microscopy revealed that understory conifers had thinner margo strands, greater maximum pore size in the margo, and more torus extensions. All of these trends may contribute to inadequate sealing of the torus. This is supported by the fact that all species showed increased vulnerability to cavitation when grown in the understory. Although evaporative demand in an understory environment is low, a rapid change into fully exposed conditions could be detrimental for shaded conifers.

Inequality of size and size increment in Pinus banksiana in relation to stand dynamics and annual growth rate.

BACKGROUND AND AIMS: Changes in size inequality in tree populations are often attributed to changes in the mode of competition over time. The mode of competition may also fluctuate annually in response to variation in growing conditions. Factors causing growth rate to vary can also influence competition processes, and thus influence how size hierarchies develop. METHODS: Detailed data obtained by tree-ring reconstruction were used to study annual changes in size and size increment inequality in several even-aged, fire-origin jack pine (Pinus banksiana) stands in the boreal shield and boreal plains ecozones in Saskatchewan and Manitoba, Canada, by using the Gini and Lorenz asymmetry coefficients. KEY RESULTS: The inequality of size was related to variables reflecting long-term stand dynamics (e.g. stand density, mean tree size and average competition, as quantified using a distance-weighted absolute size index). The inequality of size increment was greater and more variable than the inequality of size. Inequality of size increment was significantly related to annual growth rate at the stand level, and was higher when growth rate was low. Inequality of size increment was usually due primarily to large numbers of trees with low growth rates, except during years with low growth rate when it was often due to small numbers of trees with high growth rates. The amount of competition to which individual trees were subject was not strongly related to the inequality of size increment. CONCLUSIONS: Differences in growth rate among trees during years of poor growth may form the basis for development of size hierarchies on which asymmetric competition can act. A complete understanding of the dynamics of these forests requires further evaluation of the way in which factors that influence variation in annual growth rate also affect the mode of competition and the development of size hierarchies.

Carbon isotope discrimination and water stress in trembling aspen following variable retention harvesting.

Variable retention harvesting (VRH) has been proposed as a silvicultural practice to maintain biodiversity and ecosystem integrity. No previous study has examined tree carbon isotope discrimination to provide insights into water stress that could lead to dieback and mortality of trees following VRH. We measured and compared the carbon isotope ratios (delta(13)C) in stem wood of trembling aspen (Populus tremuloides Michx.) before and after VRH. Eight trees were sampled from isolated residual, edge and control (interior of unharvested stand) positions from each of seven plots in three regions (Calling Lake and Drayton Valley, Alberta and Lac Duparquet, Québec). After VRH, the general trend in mean delta(13)C was residual > edge > control trees. Although this trend is indicative of water stress in residual trees, it also suggests that edge trees received some sheltering effect, reducing their stress compared with that of residuals. A strong inverse relationship was found between the delta(13)C values and the mean annual precipitation in each region. The trend in mean delta(13)C signature was Calling Lake > Drayton Valley > Lac Duparquet trees. These results suggest that residual or edge trees in drier regions are more likely to suffer water stress following VRH. We also observed a trend of greater delta(13)C in stout trees compared with slender trees, both before and after VRH. The evidence of greater water stress in stout trees likely occurred because of a positive relationship between stem diameter and crown volume per basal area. Our results provide evidence that water stress could be the driving mechanism leading to dieback and mortality of residual trees shortly after VRH. Additionally, the results from edge trees indicate that leaving hardwood residuals in larger patches or more sheltered landscape positions could reduce the water stress to which these trees are subjected, thereby reducing dieback and mortality.

Reducing stem bending increases the height growth of tall pines.

The hypothesis was tested that upper limits to height growth in trees are the result of the increasing bending moment of trees as they grow in height. The increasing bending moment of tall trees demands increased radial growth at the expense of height growth to maintain mechanical stability. In this study, the bending moment of large lodgepole pine (Pinus contorta Dougl. Ex Loud. var. latifolia Engelm.) was reduced by tethering trees at 10 m height to counter the wind load. Average bending moment of tethered trees was reduced to 38% of control trees. Six years of tethering resulted in a 40% increase in height growth relative to the period before tethering. By contrast, control trees showed decreased height growth in the period after tethering treatment. Average radial growth along the bole, relative to height growth, was reduced in tethered trees. This strongly suggests that mechanical constraints play a crucial role in limiting the height growth of tall trees. Analysis of bending moment and basal area increment at both 10 m and 1.3 m showed that the amount of wood added to the stem was closely related to the bending moment produced at these heights, in both control and tethered trees. The tethering treatment also resulted in an increase in the proportion of latewood at the tethering height, relative to 1.3 m height. For untethered control trees, the ratio of bending stresses at 10 m versus 1.3 m height was close to 1 in both 1998 and 2003, suggesting a uniform stress distribution along the outer surface of the bole.

Carbohydrate transfer through root grafts to support shaded trees.

We investigated whether root grafts between lodgepole pine (Pinus contorta var. latifolia Dougl. ex. Loud.) trees can transfer sufficient carbohydrate reserves from a source tree to a grafted sink tree to affect the vigor of trees growing in a light-limited environment. Eleven plots were established in early spring and two grafted tree pairs and two independent non-grafted trees were selected at each plot. One tree in a grafted pair and one non-grafted tree were shaded at each plot, whereas the remaining trees were non-shaded during the experimental period. Shaded trees had significantly lower carbohydrate reserves and smaller crowns than non-shaded trees following one growing season. Grafted shaded trees had significantly higher root total nonstructural carbohydrate concentrations than non-grafted shaded trees, indicating that root grafts partially offset the effects of shading. Also, large root grafts transferred proportionately more carbohydrates to the shaded trees than small root grafts. Carbohydrates transferred through root grafts could allow grafted trees to persist under conditions where non-grafted trees would be removed by competition.

Signals controlling root suckering and adventitious shoot formation in aspen (Populus tremuloides).

We determined the effects of removal of leaves, stem axillary buds, or the entire shoot on root suckering (adventitious shoot formation by roots) and basal stem sprouts in 3- and 4-year-old potted seedlings of aspen (Populus tremuloides Michx.). The greatest number of root suckers (67.9 +/- 8.5 per plant) emerged after excision of the entire shoot. Defoliated and debudded stems were the major source of inhibitory agents for root suckering, although axillary buds and developing new leaves also exerted a significant inhibitory effect. Removal of mature leaves had only a minor effect on root suckering. Removal of a continuous band of bark (girdling) at the base of the stem consistently stimulated growth of adventitious shoots from the stem below the girdle and occasionally promoted root suckering. Exogenous application of indole-3-acetic acid to excised stumps inhibited root suckering and basal stem sprouting. Naphthylphthalamic acid (NPA), an auxin polar transport inhibitor, had no effect on root suckering or stem sprouting when it was applied to the bark of the basal stem. However, NPA significantly increased root suckering when it was applied to the exposed surface of xylem after girdling. These results suggest that polar transport of auxin in the xylem parenchyma is an important inhibitor of root suckering. On decapitated stems, vacuum extraction of xylem sap from the root system lowered the frequency of root suckering compared with decapitation alone, indicating that substance(s) originating in the root system also play a significant role in controlling root suckering.

A unified nomenclature for quantification and description of water conducting properties of sapwood xylem based on Darcy's law.

The literature dealing with the water conducting properties of sapwood xylem in trees is inconsistent in terminology, symbols and units. This has resulted from confusion in the use of either an analogy to Ohm's law or Darcy's law as the basis for nomenclature. Ohm's law describes movement of electricity through a conductor, whereas Darcy's law describes movement of a fluid (liquid or gas) through a porous medium. However, it is generally not realized that, in their full notation, these laws are mathematically equivalent. Despite this, plant physiologists have failed to agree on a convention for nomenclature. As a result, the study of water movement through sapwood xylem is confusing, especially for scientists entering the field. To improve clarity, we suggest the adoption of a single nomenclature that can be used by all plant physiologists when describing water movement in xylem. Darcy's law is an explicit hydraulic relationship and the basis for established theories that describe three-dimensional saturated and unsaturated flow in porous media. We suggest, therefore, that Darcy's law is the more appropriate theoretical framework on which to base nomenclature describing sapwood hydraulics. Our proposed nomenclature is summarized in a table that describes conventional terms, with their formulae, dimensions, units and symbols; the table also lists the many synonyms found in recent literature that describe the same concepts. Adoption of this proposal will require some changes in the use of terminology, but a common rigorous nomenclature is needed for efficient and clear communication among scientists.

Growth response and sapwood hydraulic properties of young lodgepole pine following repeated fertilization.

We examined how tree growth and hydraulic properties of branches and boles are influenced by periodic (about 6 years) and annual fertilization in two juvenile lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) stands in the interior of British Columbia, Canada. Mean basal area (BA), diameter at breast height (DBH) and height increments and percent earlywood and sapwood hydraulic parameters of branches and boles were measured 7 or 8 years after the initial treatments at Sheridan Creek and Kenneth Creek. At Sheridan Creek, fertilization significantly increased BA and DBH increments, but had no effect on height increment. At Kenneth Creek, fertilization increased BA, but fertilized trees had significantly lower height increments than control trees. Sapwood permeability was greater in lower branches of repeatedly fertilized trees than in those of control trees. Sapwood permeabilities of the lower branches of trees in the control, periodic and annual treatments were 0.24 x 10(-12), 0.35 x 10(-12) and 0.45 x 10(-12) m2 at Kenneth Creek; and 0.41 x 10(-12), 0.54 x 10(-12) and 0.65 x 10(-12) m2 at Sheridan Creek, respectively. Annual fertilization tended to increase leaf specific conductivities and Huber values of the lower branches of trees at both study sites. We conclude that, in trees fertilized annually, the higher flow capacity of lower branches may reduce the availability of water to support annual growth of the leader and upper branches.

Stem sapwood permeability in relation to crown dominance and site quality in self-thinning fire-origin lodgepole pine stands.

Stem sapwood hydraulic permeability, tree leaf area, sapwood basal area, earlywood to latewood ratio of annual rings, radial variation in hydraulic permeability and stem hydraulic capacity were examined in dominant (D), codominant (CD) and suppressed (SP) lodgepole pine (Pinus contorta Dougl. ex Loud.) trees growing on medium and poor sites. Hydraulic permeability on a sapwood area basis (ks) was lower in suppressed trees (0.71 x 10(-12) m2) compared to dominants (1.97 x 10(-12) m2) and codominants (1.79 x 10(-12) m2), and higher on medium than on poor sites. The leaf/sapwood area ratio (S) varied with crown dominance position (D > CD > SP) but not by site type. Leaf specific conductivity (kL) did not vary between crown classes or site types. The relationship between leaf area and stem hydraulic supply capacity (Q*) was strong, but differed among crown classes. Dominant trees and trees from the medium sites had a greater proportion of earlywood in outer rings of sapwood than suppressed trees. Sapwood permeability declined from the cambium to the sapwood-heartwood boundary in all samples, but the decline was more gradual in dominant trees compared to codominant and suppressed trees; differences in the radial variation in sapwood permeability may be related to differences in S. Sapwood permeability is positively related to crown dominance, whereas subdominant (CD and SP) trees have greater Q* in relation to leaf area, leading us to propose that this may give subdominant trees a survival advantage, slowing self-thinning.