Size-mediated tree transpiration along soil drainage gradients in a boreal black spruce forest wildfire chronosequence

Boreal forests are crucial to climate change predictions because of their large land area and ability to sequester and store carbon, which is controlled by water availability. Heterogeneity of these forests is predicted to increase with climate change through more frequent wildfires, warmer, longer growing seasons and potential drainage of forested wetlands. This study aims at quantifying controls over tree transpiration with drainage condition, stand age and species in a central Canadian black spruce boreal forest. Heat dissipation sensors were installed in 2007 and data were collected through 2008 on 118 trees (69 Picea mariana (Mill.) Britton, Sterns & Poggenb. (black spruce), 25 Populus tremuloides Michx. (trembling aspen), 19 Pinus banksiana Lamb. (jack pine), 3 Larix laricina (Du Roi) K. Koch (tamarack) and 2 Salix spp. (willow)) at four stand ages (18, 43, 77 and 157 years old) each containing a well- and poorly-drained stand. Transpiration estimates from sap flux were expressed per unit xylem area, J(S), per unit ground area, E(C) and per unit leaf area, E(L), using sapwood (A(S)) and leaf (A(L)) area calculated from stand- and species-specific allometry. Soil drainage differences in transpiration were variable; only the 43- and 157-year-old poorly-drained stands had?～?50% higher total stand E(C) than well-drained locations. Total stand E(C) tended to decrease with stand age after an initial increase between the 18- and 43-year-old stands. Soil drainage differences in transpiration were controlled primarily by short-term physiological drivers such as vapor pressure deficit and soil moisture whereas stand age differences were controlled by successional species shifts and changes in tree size (i.e., A(S)). Future predictions of boreal climate change must include stand age, species and soil drainage heterogeneity to avoid biased estimates of forest water loss and latent energy exchanges.     

Edge effects and forest water use: A field study in a mixed deciduous woodland

A field study was carried out in a mixed deciduous forest in order to measure the spatial variability of evapotranspiration in relation to distance from the nearest forest edge. Throughfall was collected in storage gauges in a transect across the edge. Transpiration was measured at the tree scale by means of the sap flux technique. Thermal dissipation probes were inserted into the hydro-active sapwood of 12–16 sample trees at a time covering four species. The sample trees were located close to a north- and a south-facing forest edge and between 3 and 69 m away from the nearest edge. The probes were moved to new trees about once a month and in total 71 trees were sampled. Sap flux densities were compared with potential evaporation and scaled up to the stand through multiplication with sapwood area per unit ground area. No significant edge effect on interception evaporation could be detected but there was a large influence on stand transpiration which increased towards the edge. In ash (Fraxinus excelsior L.), this increase resulted mainly from enhanced sap flux density (by 33–82%, depending on the size class) in trees located at the edge, whereas in oak (Quercus robur L.) the sap flux density was similar in edge and inner trees and an effect was only found at the stand scale in the way that the total basal area, per unit ground area, was larger near the forest edge than in the forest interior. Hawthorn (Crataegus monogyna L.) and field maple (Acer campestre L.), which occurred mainly in the understorey, were only weakly affected by the proximity to an edge. At the stand scale the total seasonal transpiration varied between 354 mm in the forest interior (>45 m away from the edge) and 565 mm at the forest edge (<15 m away from the edge), whilst the potential evaporation over the same period was 571 mm. This corresponds to Priestley–Taylor coefficients of 0.78 in the interior and 1.25 at the edge, whilst intermediate numbers were found for the area between the edge and inner zones. Using these results to calculate the average water loss per unit ground area of hypothetical woodlands of various sizes, it is shown that the edge effect dominates the water use of small forests and becomes negligible only for woodlands larger than 100 ha.     

Species-specific partitioning of soil water resources in an old-growth Douglas-fir-western hemlock forest

Although tree- and stand-level estimates of forest water use are increasingly common, relatively little is known about partitioning of soil water resources among co-occurring tree species. We studied seasonal courses of soil water utilization in a 450-year-old Pseudotsuga menziesii (Mirb.) Franco-Tsuga heterophylla (Raf.) Sarg. forest in southwestern Washington State. Soil volumetric water content (theta) was continuously monitored with frequency domain capacitance sensors installed at eight depths from 0.2 to 2 m at four locations in the vicinity of each species. Vertical profiles of root distribution and seasonal and daily courses of hydraulic redistribution (HR), sap flow and tree water status were also measured. Mean root area in the upper 60 cm of soil was significantly greater in the vicinity of T. heterophylla trees. However, seasonal water extraction on a root area basis was significantly greater near P. menziesii trees at all depths between 15 and 65 cm, leading to significantly lower water storage in the upper 65 cm of soil near P. menziesii trees at the end of the summer dry season. Greater apparent efficiency of P. menziesii roots at extracting soil water was attributable to a greater driving force for water uptake rather than to differences in root hydraulic properties between the species. The dependence of HR on theta was similar in soil near individuals of both species, but seasonal maximum rates of HR were greater in soil near P. menziesii because minimum values of theta were lower, implying a steeper water potential gradient between the upper and lower soil that acted as a driving force for water efflux from shallow roots. The results provide information on functional traits relevant for understanding the ecological distributions of these species and have implications for spatial variability of processes such as soil respiration and nutrient cycling.     

Trends in bole biomass accumulation, net primary production and tree mortality in Pseudotsuga menziesii forests of contrasting age

Although it is generally accepted that the rate of accumulation of biomass declines as forests age, little is known about the relative contributions to this decline of changes in net primary production (NPP) and tree mortality. We used 10-15 years of observations of permanent plots in three small watersheds in and near the H.J. Andrews Experimental Forest, Oregon, to examine these issues. The three watersheds are of similar elevation and potential productivity and support young (29 years at last measurement), mature (approximately 100 years) and old (approximately 400 years) forest dominated by Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg. Accumulation of tree bole biomass was greatest in the young stand, reaching approximately 7 Mg ha-1 year-1 in the last measurement interval. Bole biomass accumulation was relatively constant (approximately 4-5 Mg ha-1 year-1) in the mature stand, and there was no net accumulation of bole biomass in the old-forest stand. The NPP of boles increased with time in the young stand, from approximately 3 to approximately 7 Mg ha-1 year-1, but was nearly constant in the mature and old-forest stands, at approximately 6 and 3-4 Mg ha-1 year-1, respectively. Mortality increased slowly in the young stand (from < 0.1 to 0.3 Mg ha-1 year-1), but fluctuated between 1-2 and 2-6 Mg ha-1 year-1 in the mature and old-forest stands, respectively. Thus, declining biomass accumulation with stand age reflects, in approximately equal amounts, both decreasing NPP and increasing mortality.     

Radial variation in sap velocity as a function of stem diameter and sapwood thickness in yellow-poplar trees

Canopy transpiration and forest water use are frequently estimated as the product of sap velocity and cross-sectional sapwood area. Few studies, however, have considered whether radial variation in sap velocity and the proportion of sapwood active in water transport are significant sources of uncertainty in the extrapolation process. Therefore, radial profiles of sap velocity were examined as a function of stem diameter and sapwood thickness for yellow-poplar (Liriodendron tulipifera L.) trees growing on two adjacent watersheds in eastern Tennessee. The compensation heat pulse velocity technique was used to quantify sap velocity at four equal-area depths in 20 trees that ranged in stem diameter from 15 to 69 cm, and in sapwood thickness from 2.1 to 14.8 cm. Sap velocity was highly dependent on the depth of probe insertion into the sapwood. Rates of sap velocity were greatest for probes located in the two outer sapwood annuli (P1 and P2) and lowest for probes in closest proximity to the heartwood (P3 and P4). Relative sap velocities averaged 0.98 at P1, 0.66 at P2, 0.41 at P3 and 0.35 at P4. Tree-specific sap velocities measured at each of the four probe positions, divided by the maximum sap velocity measured (usually at P1 or P2), indicated that the fraction of sapwood functional in water transport (f(S)) varied between 0.49 and 0.96. There was no relationship between f(S) and sapwood thickness, or between f(S) and stem diameter. The fraction of functional sapwood averaged 0.66 +/- 0.13 for trees on which radial profiles were determined. No significant depth-related differences were observed for sapwood density, which averaged 469 kg m(-3) across all four probe positions. There was, however, a significant decline in sapwood water content between the two outer probe positions (1.04 versus 0.89 kg kg(-1)). This difference was not sufficient to account for the observed radial variation in sap velocity. A Monte-Carlo analysis indicated that the standard error in estimated mean f(S) declined rapidly with increasing sample size. At n = 10, the coefficient of variation in mean f(S) was 7% and at n = 15 it was slightly less than 5%. These observations indicate that radial variation in sap velocity is an important, albeit often overlooked, source of uncertainty in the scaling process. Failure to recognize that not all sapwood is functional in water transport will introduce systematic bias into estimates of both tree and stand water use. Future studies should devise sampling strategies for assessing radial variation in sap velocity and such strategies should be used to identify the magnitude of this variation in a range of non-, diffuse- and ring-porous trees.     

Transpiration and hydraulic traits of old and regrowth eucalypt forest in southwestern Australia

We tested the hypothesis that overstorey of eucalypt forest dominated by tall, large diameter trees uses less water than regrowth stands in the high rainfall zone (>1100 mm year⁻¹) of the northern jarrah (Eucalyptus marginata) forest in southwestern Australia. We measured leaf area, cover, sapwood area and sapwood density at three paired old and regrowth stands. We also measured sapflow velocity at one paired stand (Dwellingup) from June 2007 to October 2008. Old stands had more basal area but less foliage cover, less leaf area and slightly thinner sapwood. The ratio of sapwood area to basal area decreased markedly as tree size increased. Sapwood area of the regrowth forest stands (6.6 ± 0.30 m² ha⁻¹) was nearly double that of the old stands (3.4 ± 0.17 m² ha⁻¹), despite larger basal area at the old stands. Leaf area index of the regrowth stands (2.1 ± 0.26) was only one-third larger than that at the old stands (1.5 ± 0.15); hence, the ratio of leaf area to sapwood area was larger in old stands than in regrowth stands (0.45 ± 0.022 m² cm⁻² versus 0.32 ± 0.045 m² cm⁻²). Our results are consistent with theories that trees have evolved to optimize carbon gain rather than maintain stomatal conductance. Neither sapwood density (540–650 kg m⁻³) nor sap velocity differed greatly between regrowth and old stands. At the old forest site, daily transpiration rose from 0.5 mm day⁻¹ in winter to 0.9 mm day⁻¹ in spring–summer, compared to 0.9 mm day⁻¹ and 1.8 mm day⁻¹ at the regrowth site. Annual water use by the overstorey trees was estimated to be ∼230 mm year⁻¹ for the old stand and ∼500 mm year⁻¹ at the regrowth stand, or 20% and 44% of annual rainfall. The overwhelming role of stand sapwood area in determining stand water use, combined with the marked changes in the ratio of sapwood area to basal area with tree age and size, suggest that stand overstorey structure can be managed to alter overstorey water use and catchment water yield. Silviculture to promote old-forest-like attributes may be a viable means of delivering multiple water and conservation benefits.     

Forest transpiration—targeted through xylem sap flux assessment versus hydrological modeling

The assessment of forest transpiration rates is crucial for determining plant-available soil water consumption and drought risk of trees. Xylem sap flux measurements have been used increasingly to quantify stand transpiration in forest ecosystems. Here, we compare this empirical approach with hydrological modeling on the basis of a stand transpiration dataset of adult beech (Fagus sylvatica), which was acquired across Bavaria, Germany, at eight forest sites. Xylem sap flux sensors were installed in five dominant trees each. Two tree to stand upscaling approaches, related to site-specific (1) sapwood area or (2) to leaf area index, were compared. The outcome was examined each in relation to process-based stand hydrological modeling, using LWF-BROOK90. Distinct relationships between tree diameter at breast height (1.30 m) and sapwood area-weighted sap flux along the radial profile became apparent across the study sites, confirming a generic allometric basis for stand-level upscaling of transpiration. The two upscaling approaches did not differ in outcome, representatively covering stand structure for comparison with modeling. Differential analysis yielded high agreement between the empirical and modeling approaches throughout most of the study period, although LWF-BROOK90 tended to overestimate sap flux measurements under low soil moisture. The two empirical approaches proved reliable for even-aged beech stands, as performance under high stand-structural heterogeneity awaits clarification. Findings advance stand-level hydrological modeling regarding coverage of stomatal behavior during temporary limitation in water availability.     

Age-related changes in ecosystem structure and function and effects on water and carbon exchange in ponderosa pine

As forests age, their structure and productivity change, yet in some cases, annual rates of water loss remain unchanged. To identify mechanisms that might explain such observations, and to determine if widely different age classes of forests differ functionally, we examined young (Y, approximately 25 years), mature (M, approximately 90 years) and old (O, approximately 250 years) ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) stands growing in a drought-prone region of central Oregon. Although the stands differed in tree leaf area index (LAIT) (Y = 0.9, M = 2.8, O = 2.1), cumulative tree transpiration measured by sap flow did not differ substantially during the growing season (100-112 mm). Yet when water was readily available, transpiration per unit leaf area of the youngest trees was about three times that of M trees and five times that of O trees. These patterns resulted from a nearly sixfold difference in leaf specific conductance (KL) between the youngest and oldest trees. At the time of maximum transpiration in the Y stand in May-June, gross carbon uptake (gross ecosystem production, GEP) was similar for Y and O stands despite an almost twofold difference in stand leaf area index (LAIS). However, the higher rate of water use by Y trees was not sustainable in the drought-prone environment, and between spring and late summer, KL of Y trees declined fivefold compared with a nearly twofold decline for M trees and a < 30% reduction in O trees. Because the Y stand contained a significant shrub understory and more exposed soil, there was no appreciable difference in mean daily latent energy fluxes between the Y stand and the older stands as measured by the eddy-covariance technique. These patterns resulted in 60 to 85% higher seasonal GEP and 55 to 65% higher water-use efficiency at the M and O stands compared with the Y stand.     

Regulation of water flux through tropical forest canopy trees: do universal rules apply?

Tropical moist forests are notable for their richness in tree species. The presence of such a diverse tree flora presents potential problems for scaling up estimates of water use from individual trees to entire stands and for drawing generalizations about physiological regulation of water use in tropical trees. We measured sapwood area or sap flow, or both, in 27 co-occurring canopy species in a Panamanian forest to determine the extent to which relationships between tree size, sapwood area and sap flow were species-specific, or whether they were constrained by universal functional relationships between tree size, conducting xylem area, and water use. For the 24 species in which active xylem area was estimated over a range of size classes, diameter at breast height (DBH) accounted for 98% of the variation in sapwood area and 67% of the variation in sapwood depth when data for all species were combined. The DBH alone also accounted for > or = 90% of the variation in both maximum and total daily sap flux density in the outermost 2 cm of sapwood for all species taken together. Maximum sap flux density measured near the base of the tree occurred at about 1,400 h in the largest trees and 1,130 h in the smallest trees studied, and DBH accounted for 93% of the variation in the time of day at which maximum sap flow occurred. The shared relationship between tree size and time of maximum sap flow at the base of the tree suggests that a common relationship between diurnal stem water storage capacity and tree size existed. These results are consistent with a recent hypothesis that allometric scaling of plant vascular systems, and therefore water use, is universal.     

Age-related effects on leaf area/sapwood area relationships,canopy transpiration and carbon gain of Norway spruce stands (Picea abies) in the Fichtelgebirge,Germany

Stand age is an important structural determinant of canopy transpiration (E(c)) and carbon gain. Another more functional parameter of forest structure is the leaf area/sapwood area relationship, A(L)/A(S), which changes with site conditions and has been used to estimate leaf area index of forest canopies. The interpretation of age-related changes in A(L)/A(S) and the question of how A(L)/A(S) is related to forest functions are of current interest because they may help to explain forest canopy fluxes and growth. We conducted studies in mature stands of Picea abies (L.) Karst. varying in age from 40 to 140 years, in tree density from 1680 to 320 trees ha(-1), and in tree height from 15 to 30 m. Structural parameters were measured by biomass harvests of individual trees and stand biometry. We estimated E(c) from scaled-up xylem sap flux of trees, and canopy-level fluxes were predicted by a three-dimensional microclimate and gas exchange model (STANDFLUX). In contrast to pine species, A(L)/A(S) of P. abies increased with stand age from 0.26 to 0.48 m(2) cm(-2). Agreement between E(c) derived from scaled-up sap flux and modeled canopy transpiration was obtained with the same parameterization of needle physiology independent of stand age. Reduced light interception per leaf area and, as a consequence, reductions in net canopy photosynthesis (A(c)), canopy conductance (g(c)) and E(c) were predicted by the model in the older stands. Seasonal water-use efficiency (WUE = A(c)/E(c)), derived from scaled-up sap flux and stem growth as well as from model simulation, declined with increasing A(L)/A(S) and stand age. Based on the different behavior of age-related A(L)/A(S) in Norway spruce stands compared with other tree species, we conclude that WUE rather than A(L)/A(S) could represent a common age-related property of all species. We also conclude that, in addition to hydraulic limitations reducing carbon gain in old stands, a functional change in A(L)/A(S) that is related to reduced light interception per leaf area provides another potential explanation for reduced carbon gain in old stands of P. abies, even when hydraulic constraints increase in response to changes in canopy architecture and aging.     

Use of temporal patterns in vapor pressure deficit to explain spatial autocorrelation dynamics in tree transpiration

To quantify the relationship between temporal and spatial variation in tree transpiration, we measured sap flow in 129 trees with constant-heat sap flow sensors in a subalpine forest in southern Wyoming, USA. The forest stand was located along a soil water gradient from a stream side to near the top of a ridge. The stand was dominated by Pinus contorta Dougl. ex Loud. with Picea engelmannii Parry ex Engelm and Abies lasiocarpa (Hook.) Nutt. present near the stream and scattered individuals of Populus tremuloides Michx. throughout the stand. We used a cyclic sampling design that maximized spatial information with a minimum number of samples for semivariogram analyses. All species exhibited previously established responses to environmental variables in which the dominant driver was a saturating response to vapor pressure deficit (D). This response to D is predictable from tree hydraulic theory in which stomatal conductance declines as D increases to prevent excessive cavitation. The degree to which stomatal conductance declines with D is dependent on both species and individual tree physiology and increases the variability in transpiration as D increases. We quantified this variability spatially by calculating the spatial autocorrelation within 0.2-kPa D bins. Across 11 bins of D, spatial autocorrelation in individual tree transpiration was inversely correlated to D and dropped from 45 to 20 m. Spatial autocorrelation was much less for transpiration per unit leaf area and not significant for transpiration per unit sapwood area suggesting that spatial autocorrelation within a particular D bin could be explained by tree size. Future research should focus on the mechanisms underlying tree size spatial variability, and the potentially broad applicability of the inverse relationship between D and spatial autocorrelation in tree transpiration.     

Enhanced transpiration by riparian buffer trees in response to advection in a humid temperate agricultural landscape

Riparian buffers are designed as management practices to increase infiltration and reduce surface runoff and transport of sediment and nonpoint source pollutants from crop fields to adjacent streams. Achieving these ecosystem service goals depends, in part, on their ability to remove water from the soil via transpiration. In these systems, edges between crop fields and trees of the buffer systems can create advection processes, which could influence water use by trees. We conducted a field study in a riparian buffer system established in 1994 under a humid temperate climate, located in the Corn Belt region of the Midwestern U.S. (Iowa). The goals were to estimate stand level transpiration by the riparian buffer, quantify the controls on water use by the buffer system, and determine to what extent advective energy and tree position within the buffer system influence individual tree transpiration rates. We primarily focused on the water use response (determined with the Heat Ratio Method) of one of the dominant species (Acer saccharinum) and a subdominant (Juglans nigra). A few individuals of three additional species (Quercus bicolor, Betula nigra, Platanus occidentalis) were monitored over a shorter time period to assess the generality of responses. Meteorological stations were installed along a transect across the riparian buffer to determine the microclimate conditions. The differences found among individuals were attributed to differences in species sap velocities and sapwood depths, location relative to the forest edge and prevailing winds and canopy exposure and dominance. Sapflow rates for A. saccharinum trees growing at the SE edge (prevailing winds) were 39% greater than SE interior trees and 30% and 69% greater than NW interior and edge trees, respectively. No transpiration enhancement due to edge effect was detected in the subdominant J. nigra. The results were interpreted as indicative of advection effects from the surrounding crops. Further, significant differences were document in sapflow rates between the five study species, suggesting that selection of species is important for enhancing specific riparian buffer functions. However, more information is needed on water use patterns among diverse species growing under different climatic and biophysical conditions to assist policy and management decisions regarding effective buffer design.     

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.     

An analysis of sap flow in mountain ash (Eucalyptus regnans) forests of different age

Diurnal measurements of sap velocity were made in 50-, 90-, 150- and 230-year-old mountain ash (Eucalyptus regnans F. Muell.) forests in the North Maroondah catchment (southeast Australia) over the periods January 8, 1990 to April 4, 1990 and October 29, 1990 to April 16, 1991. Over the two periods, daily mean sap velocities for the four forests, in order of increasing age, were 11.5, 11.4, 9.9 and 11.8 cm h(-1) respectively. Daily mean sap velocity did not differ significantly among the 50-, 90- and 230-year-old plots. However, in the 150-year-old trees it was significantly smaller by an average of 14%. Sap velocity varied diurnally and also between positions within individual trees and among trees both within and between stands. Despite this variability, the sampling intensity and duration were sufficient to establish that behavior was highly correlated among individuals within plots. There was a significant decline with age in the overstory sapwood conducting area of these forests. In order of increasing age, the values were 6.7, 6.1, 4.2 and 4.0 m(-2) ha(-1), respectively. When combined with daily mean sap velocity, these data allowed the calculation of overstory water use. Over the experimental period, water use of the overstory decreased with age ranging, on average, from 1.86 mm day(-1) for the 50-year-old plot to 0.81 mm day(-1) for the 230-year-old plot. Mean daily water use for the two intermediate-aged forests was 1.67 and 1.00 mm day(-1), respectively. Annual water use decreased with forest age from 679 mm for the 50-year-old stand to 296 mm for the 230-year-old stand. This difference corresponds to 3.8 x 10(3) m(3) ha(-1). The annual water use of the intermediate-aged stands was 610 and 365 mm for the 90- and 150-year-old stands, respectively.     

Estimating water use of a black locust plantation by the thermal dissipation probe method in the semiarid region of Loess Plateau, China

Black locust (Robinia pseudoacacia) is a major reforestation species in the semiarid region in the Loess Plateau of China. There has been increasing concern about the sustainability of the plantations because of their possible high water-use. This study was, accordingly, undertaken to quantify the stand-scale water use of a middle-aged black locust plantation in the region. The thermal dissipation probe method was applied to 27 trees to measure sap flux densities in an experimental plot during the growing season of 2008. The monoculture stand has a basal area of 23.3 m² ha^?1 and a maximum plant area index (PAI) of 2.89. Sapwood areas were estimated by use of a regressive relationship with the diameter at breast height (DBH) for scaling up of stand transpiration. The results showed that DBH could be a good predictor of sapwood area of individual trees. The diurnal cycles of average sap flux densities differed among DBH classes. Daily transpiration can be predicted from mean daily daytime vapor pressure deficit (VPD_m) using a fitted exponential saturation model. Model variables were different among seasons, probably owing to different soil water conditions and leaf phenology. By using the derived model for each month, stand canopy transpiration over the growing season was estimated to be 73.8 mm, with an average daily value of 0.41 mm day^?1 and a maximum of 0.89 mm day^?1. The relatively small estimates of stand transpiration might be attributed to low PAI and sap wood area of the middle-aged stand.     

Dynamics of transpiration, sap flow and use of stored water in tropical forest canopy trees

In large trees, the daily onset of transpiration causes water to be withdrawn from internal storage compartments, resulting in lags between changes in transpiration and sap flow at the base of the tree. We measured time courses of sap flow, hydraulic resistance, plant water potential and stomatal resistance in co-occurring tropical forest canopy trees with trunk diameters ranging from 0.34-0.98 m, to determine how total daily water use and daily reliance on stored water scaled with size. We also examined the effects of scale and tree hydraulic properties on apparent time constants for changes in transpiration and water flow in response to fluctuating environmental variables. Time constants for water movement were estimated from whole-tree hydraulic resistance (R) and capacitance (C) using an electric circuit analogy, and from rates of change in water movement through intact trees. Total daily water use and reliance on stored water were strongly correlated with trunk diameter, independent of species. Although total daily withdrawal of water from internal storage increased with tree size, its relative contribution to the daily water budget (approximately 10%) remained constant. Net withdrawal of water from storage ceased when upper branch water potential corresponded to the sapwood water potential (Psi(sw)) at which further withdrawal of water from sapwood would have caused Psi(sw) to decline precipitously. Stomatal coordination of vapor and liquid phase resistances played a key role in limiting stored water use to a nearly constant fraction of total daily water use. Time constants for changes in transpiration, estimated as the product of whole- tree R and C, were similar among individuals (~0.53 h), indicating that R and C co-varied with tree size in an inverse manner. Similarly, time constants estimated from rates of change in crown and basal sap flux were nearly identical among individuals and therefore independent of tree size and species.     

Changes in sapwood permeability and anatomy with tree age and height in the broad-leaved evergreen species Eucalyptus regnans

Increases in plant size and structural complexity with increasing age have important implications for water flow through trees. Water supply to the crown is influenced by both the cross-sectional area and the permeability of sapwood. It has been hypothesized that hydraulic conductivity within sapwood increases with age. We investigated changes in sapwood permeability (k) and anatomy with tree age and height in the broad-leaved evergreen species Eucalyptus regnans F. Muell. Sapwood was sampled at breast height from trees ranging from 8 to 240 years old, and at three height positions on the main stem of 8-year-old trees. Variation in k was not significant among sampling height positions in young trees. However, k at breast height increased with tree age. This was related to increases in both vessel frequency and vessel diameter, resulting in a greater proportion of sapwood being occupied by vessel lumina. Sapwood hydraulic conductivity (the product of k and sapwood area) also increased with increasing tree age. However, at the stand level, there was a decrease in forest sapwood hydraulic conductivity with increasing stand age, because of a decrease in the number of trees per hectare. Across all ages, there were significant relationships between k and anatomy, with individual anatomical characteristics explaining 33-62% of the variation in k. There was also strong agreement between measured k and permeability predicted by the Hagen-Poiseuille equation. The results support the hypothesis of an increase in sapwood permeability at breast height with age. Further measurements are required to confirm this result at other height positions in older trees. The significance of tree-level changes in sapwood permeability for stand-level water relations is discussed.     

Hydrology-oriented (adaptive) silviculture in a semiarid pine plantation: How much can be modified the water cycle through forest management?

Hydrology-oriented silviculture might adapt Mediterranean forests to climatic changes, although its implementation demands a better understanding and quantification on the water fluxes. The influence of thinning intensity (high, medium, low and a control) and its effect on the mid-term (thinned plots in 1998 and 2008) on the water cycle (transpiration, soil water and interception) and growth [basal area increment (BAI)] were investigated in 55-year-old Aleppo pine trees. Thinning enhanced a lower dependence of growth on climate fluctuations. The high-intensity treatment showed significant increases in the mean annual BAI (from 4.1 to 17.3 cm²) that was maintained in the mid-term. Thinning intensity progressively increased the sap flow velocity (v _s) in all cases with respect to the control. In the mid-term, an increased functionality of the inner sapwood was also observed. Mean daily tree water use ranged from 5 (control) to 18 (high intensity) l tree^?1. However, when expressed on an area basis, daily transpiration ranged from 0.18 (medium) to 0.30 mm (control), meaning that in spite of the higher transpiration rates in the remaining trees, stand transpiration was reduced with thinning. Deep infiltration of water was also enhanced with thinning (about 30 % of rainfall) and did not compete with transpiration, as both presented opposite seasonal patterns. The changes in the stand water relationships after 10 years were well explained by the forest cover metric. The blue to green water ratio changed from 0.15 in the control to 0.72 in the high-intensity treatment, with the remaining treatments in the 0.34–0.48 range.     

Evaluation of transpiration in a Douglas-fir stand by means of sap flow measurements

Transpiration of a Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) stand was evaluated by sap flow measurements during a 4-month period. Between-tree variation in sap flow depended on crown class. On a sunny day, total transpiration was 1.6, 8.0 and 22.0 liters day(-1) for suppressed, codominant and dominant trees, respectively. Transpiration estimated by sap flow fell below potential evapotranspiration when available soil water decreased below 30% of its maximum value. Sap flow measurements gave transpiration values similar to those obtained by the water balance method.