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.     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

Persisting soil drought reduces leaf specific conductivity in Scots pine (Pinus sylvestris) and pubescent oak (Quercus pubescens)

Leaf specific conductivity (LSC; the ratio of stem conductivity (K(P)) to leaf area (A(L))), a measure of the hydraulic capacity of the stem to supply leaves with water, varies with soil water content. Empirical evidence for LSC responses to drought is ambiguous, because previously published results were subject to many confounding factors. We tested how LSC of similar-sized trees of the same population, under similar climatic conditions, responds to persistently wet or dry soil. Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) trees were compared between a dry site and a wet site in the Valais, an inner alpine valley in Switzerland. Soil water strongly influenced A(L) and K(P) and the plant components affecting K(P), such as conduit radius, conduit density and functional sapwood area. Trees at the dry site had lower LSC than trees with the same stem diameter at the wet site. Low LSC in trees at the dry site was associated with a smaller functional sapwood area and narrower conduits, resulting in a stronger reduction in K(P) than in A(L). These observations support the hypothesis that trees maintain a homeostatic water pressure gradient. An alternative hypothesis is that relatively high investments in leaves compared with sapwood contribute to carbon gain over an entire season by enabling rapid whole-plant photosynthesis during periods of high water availability (e.g., in spring, after rain events and during morning hours when leaf-to-air vapor pressure deficit is small). Dynamic data and a hydraulic plant growth model are needed to test how investments in leaves versus sapwood and roots contribute to transpiration and to maximizing carbon gain throughout entire growth seasons.     

Effect of stem height,dominance class,and site quality on sapwood permeability in loblolly pine, (Pinus taeda L.)

Sapwood permeability measurements in 30-year-old green loblolly pine (Pinus taeda L.) on the Clemson Experimental Forest in the Piedmont of South Carolina, USA demonstrated that permeability is not constant and is affected by position in the bole, distance from the pith, tree dominance class, and site. Permeability increased with increasing distance from the pith and with increasing height within the tree. Sapwood area decreased with increasing height reflecting the stem taper. Sapwood area, permeability, and conductance (permeability × sapwood area) were significantly greater on stands of higher site quality. Conductance values throughout the stem were nearly equal demonstrating how permeability and sapwood area work together to affect the trees' overall transport of water from base to crown. The significant increase in permeability and conductance associated with better sites and greater dominance class may be due to more functional tracheids in the inner sapwood. Evidently, this functional tissue enhances the trees' ability to obtain adequate nutrients and water especially on good sites.     

Constraints on physiological function associated with branch architecture and wood density in tropical forest trees

This study examined how leaf and stem functional traits related to gas exchange and water balance scale with two potential proxies for tree hydraulic architecture: the leaf area:sapwood area ratio (A(L):A(S)) and wood density (rho(w)). We studied the upper crowns of individuals of 15 tropical forest tree species at two sites in Panama with contrasting moisture regimes and forest types. Transpiration and maximum photosynthetic electron transport rate (ETR(max)) per unit leaf area declined sharply with increasing A(L):A(S), as did the ratio of ETR(max) to leaf N content, an index of photosynthetic nitrogen-use efficiency. Midday leaf water potential, bulk leaf osmotic potential at zero turgor, branch xylem specific conductivity, leaf-specific conductivity and stem and leaf capacitance all declined with increasing rho(w). At the branch scale, A(L):A(S) and total leaf N content per unit sapwood area increased with rho(w), resulting in a 30% increase in ETR(max) per unit sapwood area with a doubling of rho(w). These compensatory adjustments in A(L):A(S), N allocation and potential photosynthetic capacity at the branch level were insufficient to completely offset the increased carbon costs of producing denser wood, and exacerbated the negative impact of increasing rho(w) on branch hydraulics and leaf water status. The suite of tree functional and architectural traits studied appeared to be constrained by the hydraulic and mechanical consequences of variation in rho(w).     

华北落叶松边材透水性与叶面积空间分布的相关性的探讨

对华北落叶松林单木累积叶面积、树干边材面积、树干边材透水率及透水量的垂直分布规律及其相互关系进行了探讨。研究表明，引入树干边材面积与边材透水率的积可明显提高估测树冠叶面积垂直分布的精度，进而为精确估测林分的叶面积空间分布提供了新途径。     

Hydraulic conductance, light interception and needle nutrient concentration in Scots pine stands and their relations with net primary productivity

Aboveground xylem hydraulic conductance was determined in Scots pine (Pinus sylvestris L.) trees and stands from 7 to about 60 years of age. At the stand scale, leaf area index and net primary productivity (NPP, above- plus belowground) increased and reached a plateau at about 25-30 and 15-20 years, respectively; both parameters declined in mature stands. Stand hydraulic conductance followed a similar trend to NPP, with a maximum at about 15-20 years and a pronounced reduction in old stands. At the tree scale, annual biomass growth per unit of leaf area (growth efficiency) declined with tree age, whereas aboveground sapwood volume per unit leaf area, which is linearly related to maintenance respiration costs, steadily increased. Radiation interception per unit leaf area increased significantly with reduced leaf area index of mature stands, despite increased foliage clumping in the canopies of mature trees. Needle nutrient concentration did not change in the chronosequence. Tree hydraulic conductance per unit leaf area was strongly and positively correlated with growth efficiency. We discuss our findings in the context of growth reductions in mature and old trees, and suggest that increased hydraulic resistance and maintenance respiration costs may be the main causes of reduced carbon gain in mature and old trees.     

Derivation of stem taper from the pipe theory in a carbon balance framework

A dynamic tree growth model is described. The model derives the development of stem taper and vertical distribution of branch basal area from the pipe model, assuming that reuse of active pipes is regulated by foliage dynamics in a vertically explicit crown with a foliage distribution of constant shape. Based on empirical findings, the pipe model was modified slightly to allow the foliage/sapwood ratio to vary as a function of distance from the treetop. Growth was derived from carbon balance in a stand of different size trees that may shade each other. The model was applied to old and middle-aged trees growing in dense and sparse stands of Scots pine for which stand-level measurements are available as a chronosequence, but individual trees have been measured only once. Measured trees were compared with corresponding simulated trees for stem taper and vertical distribution of branch basal area. The results indicated that the pipe model assumptions, combined with a model of tree growth, are capable of producing realistic predictions of the vertical distribution of stem and branch diameter in trees of different sizes in the stand. A comparison of the results with a simple form of the uniform stress theory showed good agreement between the two models. However, a significant difference was found between the measured relative contribution of heartwood to total stem diameter and the predicted share of disused pipes in the stem. A possible explanation for this discrepancy is that the transition from sapwood to heartwood is gradual rather than abrupt as assumed in the model. A modification of the pipe model to incorporate a gradual transition is outlined.     

Temporal dynamics of stem expansion and contraction in savanna trees: withdrawal and recharge of stored water

Relationships between diel changes in stem expansion and contraction and discharge and refilling of stem water storage tissues were studied in six dominant Neotropical savanna (cerrado) tree species from central Brazil. Two stem tissues were studied, the active xylem or sapwood and the living tissues located between the cambium and the cork, made up predominantly of parenchyma cells (outer parenchyma). Outer parenchyma and sapwood density ranged from 320 to 410 kg m(-3) and from 420 to 620 kg m(-3), respectively, depending on the species. The denser sapwood tissues exhibited smaller relative changes in cross-sectional area per unit change in water potential compared with the outer parenchyma. Despite undergoing smaller relative changes in cross-sectional area, the sapwood released about 3.5 times as much stored water for a given change in area as the outer parenchyma. Cross-sectional area decreased earlier in the morning in the outer parenchyma than in the sapwood with lag times up to 30 min for most species. The relatively small lag time between dimensional changes of the two tissues suggested that they were hydraulically well connected. The initial morning increase in basal sap flow lagged about 10 to 130 min behind that of branch sap flow. Species-specific lag times between morning declines in branch and main stem cross-sectional area were a function of relative stem water storage capacity, which ranged from 16 to 31% of total diurnal water loss. Reliance on stored water to temporarily replace transpirational losses is one of the homeostatic mechanisms that constrain the magnitude of leaf water deficits in cerrado trees.     

Differences in delta13C and diameter growth among remnant Scots pine populations in Scotland

Published data suggest that differences in wood cellulose carbon isotope composition (delta13C) and xylem ring width among natural populations of Scots pine in Scotland (Pinus sylvestris L.) are attributable to the persistence of palaeotypes of various post-glacial migratory origins. We assessed differences in wood cellulose delta13C and ring width among Scottish Scots pine populations grown in a clone bank and in natural stands at various locations in northern and central Scotland. Ring width and wood cellulose delta13C varied significantly among natural stands. Potential water deficit was positively correlated with wood cellulose delta13C and xylem ring width in the natural stands. Neither wood cellulose delta13C nor xylem ring width of clone bank trees correlated with any climate variables at the sites from which the trees originated, indicating little adaptation to climate for these traits. Xylem ring width showed a site x population interaction for the growth sites (i.e., natural stands versus clone bank), but wood cellulose delta13C did not. These results suggest that climate variation in Scotland has not resulted in significant genetic variation in wood cellulose delta13C or xylem ring width in post-glacial populations.     

Effects of drought stress and high density stem inoculations with Leptographium wingfieldii on hydraulic properties of young Scots pine trees

We examined drought-induced changes in susceptibility of potted Scots pine (Pinus sylvestris L.) trees to a bark-beetle associated fungus (Leptographium wingfieldii Morelet, from the bark beetle Tomicus piniperda L.). Five-year-old field-grown trees were transplanted to 50-l pots and grown for 1 year before the treatments were applied. Trees in the drought treatment were subjected to several successive, 3-week-long drought cycles, with predawn water potential dropping below -2 MPa at peak drought intensity. The experimental drought cycles were more severe than the natural drought episodes usually recorded in Scots pine stands. Trees were then mass-inoculated with L. wingfieldii at a density close to the critical threshold density of inoculations (400 m(-2)) above which tree resistance is overcome. Inoculation of well-watered trees resulted in induced reaction zones around the inoculation points and very limited damage (resinosis) in the sapwood. Drought alone had no long-lasting consequences on tree water relations, except for a decrease in hydraulic conductance in the youngest segments of the main stem. However, the combination of mass-inoculation and drought stress after inoculation resulted in a dramatic loss of stem hydraulic conductivity that was paralleled by conspicuous damage to the sapwood (resinosis, drying and blue staining). There was a close correlation between amount of visible sapwood damage and loss of hydraulic conductivity. The intensity of induced reactions in the phloem was unaffected by drought stress. We conclude that tree defence against L. wingfieldii is decreased during severe drought stress, resulting in changes in the spread and action of the fungus in the sapwood but not in the phloem.     

Sapwood Basal Area and Needle Mass of Scots Pine (Pinus sylvestris L.) Trees in Central Sweden

The relationship between foliage mass or area and sapwood basalarea is of potential interest in the quest for cheap and simplemethods of estimating foliage biomass. The sapwood basal area and the needle dry weight were obtainedby destructive sampling for 153 Scots pine trees (Pinus sylvestrisL.) from 16 stands in central Sweden. The regression relationshipbetween needle mass and sapwood basal area was strong for singlestands, but it did not seem appropriate to aggregate the materialinto one overall regression, having sapwood basal area as theonly independent variable. It was shown that the needle biomassper unit of sapwood basal area varied with mean annual ringwidth in the sapwood. This information was used in an improvedoverall model.     

Butt-rot of Scots pine in Thetford Forest caused by Heterobasidion annosum: a local phenomenon

Heterobasidion annosum was found to be the main cause of decay in 50–60-year-old Scots pine in Thetford Forest in East England. On some sites, up to 15% of the trees could be suffering from H. annosum rot, with decay occasionally extending 2 m or more up the stem. Prior to this study, Scots pine had been considered as resistant to H. annosum butt-rot in Britain, except in overmature trees.     

Fine root biomass in relation to site and stand characteristics in Norway spruce and Scots pine stands

Variations in fine root biomass of trees and understory in 16 stands throughout Finland were examined and relationships to site and stand characteristics determined. Norway spruce fine root biomass varied between 184 and 370 g m(-2), and that of Scots pine ranged between 149 and 386 g m(-2). In northern Finland, understory roots and rhizomes (< 2 mm diameter) accounted for up to 50% of the stand total fine root biomass. Therefore, the fine root biomass of trees plus understory was larger in northern Finland in stands of both tree species, resulting in a negative relationship between fine root biomass and the temperature sum and a positive relationship between fine root biomass and the carbon:nitrogen ratio of the soil organic layer. The foliage:fine root ratio varied between 2.1 and 6.4 for Norway spruce and between 0.8 and 2.2 for Scots pine. The ratio decreased for both Norway spruce and Scots pine from south to north, as well as from fertile to more infertile site types. The foliage:fine root ratio of Norway spruce was related to basal area and stem surface area. The strong positive correlations of these three parameters with fine root nitrogen concentration implies that more fine roots are needed to maintain a certain amount of foliage when nutrient availability is low. No significant relationships were found between stand parameters and fine root biomass at the stand level, but the relationships considerably improved when both fine root biomass and stand parameters were calculated for the mean tree in the stand. When the northern and southern sites were analyzed separately, fine root biomass per tree of both species was significantly correlated with basal area and stem surface area per tree. Basal area, stem surface area and stand density can be estimated accurately and easily. Thus, our results may have value in predicting fine root biomass at the tree and stand level in boreal Norway spruce and Scots pine forests.     

Variation in treatability of Scots pine (Pinus sylvestris) by the chemical modification agent furfuryl alcohol dissolved in water

Scots pine (Pinus sylvestris) sapwood was investigated for variation in treatability using the wood modifying agent, furfuryl alcohol (FA) in water. The variation in treatability within trees, between trees and between different stands of Scots pine was studied. Investigated variables that reduced the residual variance significantly were: site location, latitude of site, height of trees, annual ring width, vertical and horizontal position in the tree and method of drying. Linear mixed model statistics were used and tree number was handled as a random variable. The best model reduced the treatability residual variance by 67%. Location was the single factor affecting treatability most. Differences in latitude between locations may be the reason for that. Latitude correlated negatively with the treatability. Within the trees, the treatability of sapwood increased with distance from ground and with distance from heartwood border. A small, but significantly better treatability was found for kiln dried wood (60°C) compared to air dried wood (20°C).     

Stored water use and transpiration in Scots pine: a modeling analysis with ANAFORE

We estimated daily use of stored water by Scots pine (Pinus sylvestris L.) trees growing in a temperate climate with the ANAFORE model (ANAlysis of FORest Ecosystems) and compared the simulation results with sap flow measurements. The original model was expanded with a dynamic water flow and storage model that simulates sap flow dynamics in an individual tree. ANAFORE was able to accurately simulate diurnal patterns of measured sap flow under microclimatic conditions that differ from those of the calibration period. Strong relationships were found between stored water use and several tree characteristics (diameter at breast height, sapwood area, leaf area), but not with tree height. Relative to transpiration, stored water use varied over time (between < 1% and 44% of daily transpiration). On days when transpiration was high, trees were more dependent on stored water, indicating that the contribution of internal water to transpiration is not a constant in the water budget of trees.     

Relationships between stem diameter, sapwood area, leaf area and transpiration in a young mountain ash forest

We examined relationships between stem diameter, sapwood area, leaf area and transpiration in a 15-year-old mountain ash (Eucalyptus regnans F. Muell.) forest containing silver wattle (Acacia dealbata Link.) as a suppressed overstory species and mountain hickory (Acacia frigescens J.H. Willis) as an understory species. Stem diameter explained 93% of the variation in leaf area, 96% of the variation in sapwood area and 88% of the variation in mean daily spring transpiration in 19 mountain ash trees. In seven silver wattle trees, stem diameter explained 87% of the variation in sapwood area but was a poor predictor of the other variables. When transpiration measurements from individual trees were scaled up to a plot basis, using stem diameter values for 164 mountain ash trees and 124 silver wattle trees, mean daily spring transpiration rates of the two species were 2.3 and 0.6 mm day(-1), respectively. The leaf area index of the plot was estimated directly by destructive sampling, and indirectly with an LAI-2000 plant canopy analyzer and by hemispherical canopy photography. All three methods gave similar results.     

Restoration thinning and influence of tree size and leaf area to sapwood area ratio on water relations of Pinus ponderosa

Ponderosa pine (Pinus ponderosa Dougl. ex P. Laws) forest stand density has increased significantly over the last century (Covington et al. 1997). To understand the effect of increased intraspecific competition, tree size (height and diameter at breast height (DBH)) and leaf area to sapwood area ratio (A(L):A(S)) on water relations, we compared hydraulic conductance from soil to leaf (kl) and transpiration per unit leaf area (Q(L)) of ponderosa pine trees in an unthinned plot to trees in a thinned plot in the first and second years after thinning in a dense Arizona forest. We calculated kl and Q(L) based on whole- tree sap flux measured with heat dissipation sensors. Thinning increased tree predawn water potential within two weeks of treatment. Effects of thinning on kl and Q(L) depended on DBH, A(L):A(S) and drought severity. During severe drought in the first growing season after thinning, kl and Q(L) of trees with low A(L):A(S) (160-250 mm DBH; 9-11 m height) were lower in the thinned plot than the unthinned plot, suggesting a reduction in stomatal conductance (g(s)) or reduced sapwood specific conductivity (K(S)), or both, in response to thinning. In contrast kl and Q(L) were similar in the thinned plot and unthinned plot for trees with high A(L):A(S) (260-360 mm DBH; 13-16 m height). During non-drought periods, kl and Q(L) were greater in the thinned plot than in the unthinned plot for all but the largest trees. Contrary to previous studies of ponderosa pine, A(L):A(S) was positively correlated with tree height and DBH. Furthermore, kl and Q(L) showed a weak negative correlation with tree height and a strong negative correlation with A(S) and thus A(L):A(S) in both the thinned and unthinned plots, suggesting that trees with high A(L):A(S) had lower g(s). Our results highlight the important influence of stand competitive environment on tree-size-related variation in A(L):A(S) and the roles of A(L):A(S) and drought on whole-tree water relations in response to thinning.     

Correlations between stable carbon-isotope abundance and hydraulic conductivity in Douglas-fir across a climate gradient in Oregon, USA

Stomatal conductance in trees is related to both foliar carbon-isotope abundance and stem hydraulic properties. By combining these relationships, I hypothesized that carbon-isotope abundance in foliage should vary with limitations to water movement through supporting branches. I sampled Douglas-fir branches (Pseudotsuga menziesii (Mirb.) Franco) from six sites across a climate gradient in Oregon, USA for foliar carbon-isotope abundance and stem hydraulic properties. I used a forest growth model to quantify climate-induced stomatal limitations, expressed as reduced potential transpiration, across the gradient. Foliar stable carbon-isotope abundance showed a strong inverse relationship with branch specific conductivity (hydraulic conductivity per unit functional sapwood area) and leaf-specific conductivity (hydraulic conductivity per unit leaf area). Foliar stable carbon-isotope abundance was correlated with modeled reductions in potential transpiration; however, the inclusion of leaf-specific conductivity improved the correlation by more than 30%. Combined, leaf-specific conductivity and climate-induced stomatal constraints explained 84% of the variation in foliar isotope abundance in 1994 foliage. This model was confirmed on foliage classes 1990-1993.     

Dry season conditions determine wet season water use in the wet-tropical savannas of northern Australia

Daily and seasonal patterns of transpiration were measured in evergreen eucalypt trees growing at a wet (Darwin), intermediate (Katherine) and dry site (Newcastle Waters) along a steep rainfall gradient in a north Australian savanna. Relationships between tree size and tree water use were also determined. Diameter at breast height (DBH) was an excellent predictor of sapwood area in the five eucalypt species sampled along the rainfall gradient. A single relationship existed for all species at all sites. Mean daily water use was also correlated to DBH in both wet and dry seasons. There were no significant differences in the relationship between DBH and tree water use at Darwin or Katherine. Among the sites, tree water use was lowest at Newcastle Waters at all DBHs. The relationship between DBH and tree leaf area was similar between species and locations, but the slope of the relationship was less at the end of the dry season than at the end of the wet season at all locations. There was a strong relationship between sapwood area and leaf area that was similar at all sites along the gradient. Transpiration rates were significantly lower in trees at the driest site than at the other sites, but there were no significant differences in transpiration rates between trees growing at Darwin and Katherine. Transpiration rates did not vary significantly between seasons at any site. At all sites, there was only a 10% decline in water use per tree between the wet and dry seasons. A monthly aridity index (pan evaporation/rainfall) and predawn leaf water potential showed strong seasonal patterns. It is proposed that dry season conditions exert control on tree water use during the wet season, possibly through an effect on xylem structure.