Spatial variation in sapwood area to leaf area ratio and specific leaf area within a crown of silver birch

Spatial variation in sapwood area to leaf area ratio (Huber value, HV) and specific leaf area (SLA) was examined in branches of closed-canopy trees of silver birch (Betula pendula Roth). HV increased basipetally within a crown and decreased with increasing branch order, but exhibited no significant radial trend along a primary branch. HV was primarily determined by branch position in a crown and branch diameter at the sampling point, being independent of the size of the tree and branch. Greater HV in the lower-crown branches is considered a means to mitigate differences in hydraulic transport capacity between the branches located in different canopy layers. Beside branch position and sampling location on a branch, SLA depended significantly on several other variables characterising tree and branch size. SLA increased basipetally within a crown and along a primary branch, but exhibited no significant trend with branch orders. Because height caused leaf area (A_L) to diminish more rapidly than leaf dry weight, A_L primarily determined the vertical variation in SLA.     

Tradeoffs between hydraulic efficiency and mechanical strength in the stems of four co-occurring species of chaparral shrubs

Possible tradeoffs between efficiency of water transport and mechanical strength were examined in stems of two congeneric pairs of co-occurring chaparral shrubs. First, since previously published results indicated that Adenostoma sparsifolium (Rosaceae) had greater specific conductivity (k _s or hydraulic conductivity per xylem transverse area) than A. fasciculatum, it was hypothesized that A. sparsifolium would have greater vessel lumen area per square millimeter of xylem area, and less mechanical strength, than A. fasciculatum. Secondly, since Ceanothus megacarpus (Rhamnaceae) is a non-sprouter (unable to sprout from the root crown following fire or other major disturbance) whereas C. spinosus is a sprouter and thus able to form new stems following disturbance, it was hypothesized that C. megacarpus would have greater mechanical strength, but lower k _s, than C. spinosus. Both hypotheses were supported. Based upon computer-aided image analyses, A. sparsifolum had significantly higher mean and maximum vessel diameters (16.4, 40.5 vs. 14.6, 35.7 μm), a 34% greater percent vessel lumen area, and a two-fold greater measured and theoretical k _s than A. fasciculatum. This corresponded to 14% lower stem density (wet weight/volume) and less mechanical strength, with a 37% lower modulus of elasticity (MOE) and a 30% lower modulus of rupture (MOR) than A. fasciculatum. Similarly, C.␣spinosus had a significantly higher maximum vessel diameter (52.7 vs. 41.8 μm) and a 92% higher theoretical k _s (and 43% higher measured k _s) than C. megacarpus. This corresponded to a 9% lower stem density and 20% lower MOR than for C. megacarpus. Thus, at least within these two congeneric pairs of chaparral shrubs growing together in the same habitat, there may be tradeoffs between mechanical strength and conductive efficiency of the stem xylem which correspond to differences in transport physiology and life history traits of sprouter versus non-sprouter species.     

Resistance to water flow in xylem of Picea abies (L.) Karst. trees grown under contrasting light conditions

Summary The relative hydraulic conductivity (k) of xylem and resistance (R) to water flow through trunk, primary roots and branches in Picea abies trees growing under contrasting light conditions were investigated. The xylem permeability to water was measured by forcing 10 mM water solution of KC1 through excised wood specimens. From the values of k, the sapwood transverse area and the length of conducting segments, R of the whole trunk, branches and roots was calculated. The relative conductivity of xylem in open-grown trees exceeded that of shade-grown trees by 1.4–3.1 times, while k was closely correlated with the hydraulically effective radius (R _e) of the largest tracheids (R ² was 0.85–0.94 for open- and 0.51–0.79 for shade-grown trees). Because of both a low k and a smaller sapwood area in shade-grown trees the resistance to water movement through their trunk, roots and branches was many times higher. The distribution of R between single segments of the water-conducting pathway differed considerably in trees from different sites. At high water status the largest share of the total resistance in open- as well as shade-grown trees resides in the apical part of the trunk. The contribution of the branches to total xylem resistance is supposed to increase with developing water deficit.     

Effects of branch position on water relations and gas exchange of European larch trees in an alpine community

Water relations and gas exchange were studied in the crowns of small European larch (Larix decidua Mill.) trees with respect to branch position. The upper-crown branches showed significantly higher branch sap flux rate (F _la) and branch conductance (g _b) compared to the lower crown (P<0.001). Values of leaf conductance (g _l), transpiration rate (E) and net photosynthesis (A), averaged for different ranges of atmospheric vapour pressure deficit (VPD), were also higher in the upper crown position. We suppose that the up to 2.6-fold smaller soil-to-leaf hydraulic conductance observed in the lower branches (P<0.001, compared to upper branches) could contribute to the decreased values of F _la, g _b, g _l, and E in the lower crown position. Variation in tracheid lumen diameter with respect to crown position (P<0.001) supported the hypothesis that branches growing at the crown base are hydraulically more constrained than branches located at the top of the tree. Leaf area to sapwood area ratio (A _la/A _sa) exhibited 1.4 times smaller values in lower crown (P<0.01), however, this could not compensate the effect of decreased hydraulic conductivity of the lower-crown branches.     

Control of transpiration in three coffee cultivars: the role of hydraulic and crown architecture

Water use and hydraulic architecture were studied in the coffee (Coffea arabica) cultivars San Ramon, Yellow Caturra and Typica growing in the field under similar environmental conditions. The cultivars differed in growth habit, crown architecture, basal sapwood area and total leaf surface area. Transpiration per unit leaf area (E), stomatal conductance (g _s), crown conductance (g _c), total hydraulic conductance of the soil/leaf pathway (G _t) and the stomatal decoupling coefficient, omega (Ω) (Jarvis and McNaughton 1986) were assessed over a range of soil moisture and during partial defoliation treatments. The relationship between sap flow and sapwood area was linear and appeared to be similar for the three cultivars. Variation in g _c, E, and G _t of intact plants and leaf area-specific hydraulic conductivity (k _l) of excised lateral branches was negatively correlated with variation in the ratio of leaf area to sapwood area. Transpiration, g _c, and g _s were positively correlated with G _t. Transpiration and G _t varied with total leaf area and were greatest at intermediate values (10 m²) of leaf area. Omega was greatest in Yellow Caturra, the cultivar with the greatest leaf area and a dense crown, and was smallest in Typica, the cultivar with an open crown. Differences in omega were attributable primarily to differences in leaf boundary layer conductance among the cultivars. Plants of each cultivar that were 40% defoliated maintained sap flows comparable to pretreatment plants, but expected compensatory increases in g _s were not consistently observed. Despite their contrasting crown morphologies and hydraulic architecture, the three cultivars shared common relationships between water use and hydraulic architectural traits. Received: 17 February 1999 / Accepted: 28 July 1999     

Regulation of water flux through trunks, branches, and leaves in trees of a lowland tropical forest

We studied regulation of whole-tree water use in individuals of five diverse canopy tree species growing in a Panamanian seasonal forest. A construction crane equipped with a gondola was used to access the upper crowns and points along the branches and trunks of the study trees for making concurrent measurements of sap flow at the whole-tree and branch levels, and vapor phase conductances and water status at the leaf level. These measurements were integrated to assess physiological regulation of water use from the whole-tree to the single-leaf scale. Whole-tree water use ranged from 379 kg day⁻¹ in a 35 m-tall Anacardium excelsum tree to 46 kg day⁻¹ in an 18 m-tall Cecropia longipes tree. The dependence of whole-tree and branch sap velocity and sap flow on sapwood area was essentially identical in the five trees studied. However, large differences in transpiration per unit leaf area (E) among individuals and among branches on the same individual were observed. These differences were substantially reduced when E was normalized by the corresponding branch leaf area:sapwood area ratio (LA/SA). Variation in stomatal conductance (g _s) and crown conductance (g _c), a total vapor phase conductance that includes stomatal and boundary layer components, was closely associated with variation in the leaf area-specific total hydraulic conductance of the soil/leaf pathway (G _t). Vapor phase conductance in all five trees responded similarly to variation in G _t. Large diurnal variations in G _t were associated with diurnal variation in exchange of water between the transpiration stream and internal stem storage compartments. Differences in stomatal regulation of transpiration on a leaf area basis appeared to be governed largely by tree size and hydraulic architectural features rather than physiological differences in the responsiveness of stomata. We suggest that reliance on measurements gathered at a single scale or inadequate range of scale may result in misleading conclusions concerning physiological differences in regulation of transpiration. Received: 1 October 1997 / Accepted: 6 March 1998     

Xylem structure and water transport in a twiner,a scrambler,and a shrub of Lonicera (Caprifoliaceae)

Summary Wood structure and function was investigated in different growth forms of temperate honeysuckles (Lonicera spp.). All three species had many narrow vessels and relatively few wide ones, with the measured K _h (flow rate/pressure gradient) approximately 24–55% of the theoretical K _h predicted by Poiseuille's law. Only the twiner, Lonicera japonica, had some vessels greater than 50 m in diameter. The twiner also had the narrowest stem xylem diameters, suggesting the greater maximum vessel diameter hydraulically compensated for narrow stems. Conversely, the free-standing shrub, L. maackii, had the greatest annual increments of xylem but the least percent conductive xylem implying that a great portion of the wood was involved with mechanical support. The scrambler, L, sempervirens had low maximum vessel diameter, high Huber values (= xylem area/leaf area), and low specific conductivities (= measured K _h/xylem area), much like the shrub. The greatest vessel frequency occurred in the scrambler (901 vessels · mm^-2), the highest thus far recorded in vines. The lowest Huber value and highest specific conductivity occurred in the twiner, suggesting little self-support but relatively efficient water conduction. LSC (= measured K _h/leaf area) and maximum vessel diameter of Lonicera vines were near the low end of the range for vines in general.     

The impact of vessel size on vulnerability curves: data and models for within‐species variability in saplings of aspen,Populus tremuloides Michx

The objective of this study was to quantify the relationship between vulnerability to cavitation and vessel diameter within a species. We measured vulnerability curves (VCs: percentage loss hydraulic conductivity versus tension) in aspen stems and measured vessel‐size distributions. Measurements were done on seed‐grown, 4‐month‐old aspen (Populus tremuloides Michx) grown in a greenhouse. VCs of stem segments were measured using a centrifuge technique and by a staining technique that allowed a VC to be constructed based on vessel diameter size‐classes (D). Vessel‐based VCs were also fitted to Weibull cumulative distribution functions (CDF), which provided best‐fit values of Weibull CDF constants (c and b) and P₅₀ = the tension causing 50% loss of hydraulic conductivity. We show that P₅₀ = 6.166D^?0.3134 (R² = 0.995) and that b and 1/c are both linear functions of D with R² > 0.95. The results are discussed in terms of models of VCs based on vessel D size‐classes and in terms of concepts such as the ‘pit area hypothesis’ and vessel pathway redundancy.     

Hydraulic properties and photosynthetic rates in co-occurring lianas and trees in a seasonal tropical rainforest in southwestern China

In this study, we examined wood anatomy, hydraulic properties, photosynthetic rate, and water status and osmotic regulation in three liana species and three tree species co-occurring in a seasonal tropical rain forest. Our results showed that the three liana species had larger vessel diameter, lower sapwood density, and consequently higher branch sapwood specific hydraulic conductivity (K _S) than the three tree species. Across species, K _S was positively correlated with leaf nitrogen concentration and maximum net CO₂ assimilation rate. However, it was also positively correlated with xylem water potential at 50% loss of hydraulic conductivity, indicating a trade-off between hydraulic efficiency and safety. Compared to the tree species, the liana species had higher predawn leaf water potential and lower osmotic adjustment in the dry season. The combination of more efficient water transport, higher photosynthetic rates, and their ability to access to more reliable water source at deeper soil layers in the dry season in the lianas should contribute to their fast growth.     

A plumber's-eye view of xylem water transport in woody plants

We present a practical for university-level students aimed at measuring and comparing xylem hydraulic properties of co-existing plant species. After sampling branches of several woody species in the field, their main hydraulic properties were measured using a simple set-up. Hydraulic conductivity (K_h ) was calculated as the ratio between water flow through a plant segment and the pressure gradient driving the flow. The percent reduction in conductivity due to xylem embolism (i.e. air-filled conduits) was estimated by comparing K_h before and after flushing the measure segments to remove all native embolism. Raw hydraulic conductivity was standardised by cross-sectional wood area or supported leaf area to obtain more meaningful measures of conducting capacity. The results showed differences among study species, particularly between conifers and angiosperms. These differences are briefly discussed in terms of wood anatomy and the general biology of the species. Overall the practical provides a good opportunity for students to appreciate the main aspects of xylem water transport and the constraints it imposes on plant water relations.     

Hydraulic Parameters Measured in 1-Year-Old Twigs of some Mediterranean Species with Diffuse-Porous Wood: Changes in Hydraulic Conductivity and Their Possible Functional Significance

We report measurements of hydraulic conductivity of Vitis oinifera,Oleo europaea and Populus deltoides 1-year-old twigs. Singleserial internodes were tested for the volume flow rate whichwas related to: (a) the xylem tissue cross-sectional area, (b)the vessel lumina cross-sectional area and (c) the leaf surfacearea supplied by a given stem section. From this, whole xylemhydraulic conductivity (L_x), vessel lumina hydraulic conductivity(L_xv) and leaf specific conductivity (LSC) were calculated.All the three parameters turned out to be linearly related toeach other. This is kcause: (a) the leaf surface area (A₁) waslinearly related to xylem cross-sectional area (A_x and (b) theratio of the vessel lumina cross-sectional area (A_xv) to xylemcross-sectional area (A_x) was approximately constant along thetwigs. Moreover, the hydraulic conductivity of twig segmentswhere buds grow most actively (distal internodes in V. viniferaand proximal ones in O. europaea) was much lower than in therest of the twigs. A possible role played by these ‘constricted’twig regions is discussed. Key words: Changes, Hydraulic conductivity, Stem     

The hydraulic architecture of balsam fir (Abies balsamea)

Leaf-specific conductivities (LSCs – hydraulic conductivity per dry weight of supplied leaves). Huber values (transverse sapwood area per dry weight of supplied leaves), specific conductivity (hydraulic conductivity per transverse sapwood area) and tracheid diameters were measured throughout the trunk and crown of 20-year-old trees of Abies balsamca (L.) Mill. Measured specific conductivity was proportional to the radius to the fourth power of tracheids. LSCs, which indicate the relative water availability to different plant parts, are much higher in the trunk than in first order branches, and lowest in second order branches. The structural basis for this "hydraulic hierarchy" lies both in Huber values and in tracheid diameters. For similar diameter stem segments, there was no statistically significant difference for trunks versus branches in specific conductivity. However, in old parts of the tree, trunks are wider than supported branches and producer wider tracheids resulting in greater specific conductivities than in branches. In vigorous trees with strong apical control, Huber values were 12.0 times greater in the trunk than in similar diameter branch segments. In slow-growing trees with weak apical control, Huber values were 2.2 times greater in the trunk versus similar branch segments.     

Single‐vessel flow measurements indicate scalariform perforation plates confer higher flow resistance than previously estimated

During vessel evolution in angiosperms, scalariform perforation plates with many slit‐like openings transformed into simple plates with a single circular opening. The transition is hypothesized to have resulted from selection for decreased hydraulic resistance. Previously, additional resistivity of scalariform plates was estimated to be small – generally 10% or less above lumen resistivity – based on numerical and physical models. Here, using the single‐vessel technique, we directly measured the hydraulic resistance of individual xylem vessels. The resistivity of simple‐plated lumens was not significantly different from the Hagen–Poiseuille (HP) prediction (+6 ± 3.3% mean deviation). In the 13 scalariform‐plated species measured, plate resistivity averaged 99 ± 13.7% higher than HP lumen resistivity. Scalariform species also showed higher resistivity than simple species at the whole vessel (+340%) and sapwood (+580%) levels. The strongest predictor of scalariform plate resistance was vessel diameter (r² = 0.84), followed by plate angle (r² = 0.60). An equation based on laminar flow through periodic slits predicted single‐vessel measurements reasonably well (r² = 0.79) and indicated that Baileyan trends in scalariform plate evolution maintain an approximate balance between lumen and plate resistances. In summary, we found scalariform plates of diverse morphology essentially double lumen flow resistance, impeding xylem flow much more than previously estimated.     

Axial and radial profiles in conductivities, water storage and native embolism in trunks of young and old-growth ponderosa pine trees

Ponderosa pine has very wide sapwood, and yet the spatial and temporal use of that sapwood for water transport is poorly understood. Moreover, there have been few comparisons of function in tips of old-growth trees in comparison with young trees. In the present study, axial and radial specific conductivity (k_s), leaf specific conductivity (LSC), leaf specific conductance (k_l), native embolism and the compartmentalization of sapwood water storage were characterized in trunks of young and old-growth trees. Trunks of young trees had lower k_s, lower LSC and lower native embolism [corresponding to 5% loss of conductivity (PLC)] than trunks of old-growth trees. However, k_l in young trees was 3.5 times higher than in old-growth trees, supporting the hypothesis that tall trees have a reduced ability to transport water to their leaves. Water storage (capacitance) of young trees was not significantly different than at the base of old-growth trees. Although the top of the old-growth trees had similar k_s, LSC and k_l to the young trees for a given cambial age, they had higher native embolism and lower capacitance. There was no trade-off between k_s and native embolism at any height. In the tree crown, outer sapwood had 35–50% higher k_s than the inner sapwood and 17–25 PLC lower native embolism. At the base of the old trees, there was no significant difference in native embolism between the outer, middle and inner sapwood, showing that refilling of embolisms was complete despite the 130-year difference in wood age among these radial positions. Although during the dry season the inner sapwood tended to be more saturated than the outer sapwood, the outer part of the sapwood contributed up to 60% of the overall stored water. Safer xylem, higher capacitance and higher k_l would appear adaptive in the young trees for regulating their water resource, which is likely to be less reliable than the water availability of older trees with their more developed root system.     

The influence of drought on the relationship between leaf and conducting sapwood area in Eucalyptus globulus and Eucalyptus nitens

 Development of the relationship between leaf area (A _l ) and sapwood area (A _s ) was investigated in two important hardwoods, Eucalyptus globulus (Labill) and E. nitens (Deane and Maiden) Maiden, growing in an experimental plantation established in a low rainfall zone (approx. 515 mm year^–1) of Tasmania. The experiment compared irrigated controls and a rainfed treatment which was subjected to cyclical summer droughts from age 1 to 6 years old. Leaf area and sapwood area were determined by destructive sampling at ages 2, 3 and 6 years old. There was no effect of stand age on A _l :A _s when sapwood area was measured at crown break. At age 3 years old A _l :A _s was significantly greater in the rainfed than the irrigated trees. It was concluded that this difference was due to earlier canopy closure in the irrigated trees. When the plantation was 6 years old A _l :A _s was significantly greater in the irrigated than the rainfed treatment. An analysis based on an equation which links A _l :A _s with transpiration and volumetric flow rate (Whitehead et al. 1984) was used to infer a positive correlation between stem hydraulic conductivity (k _h ) and water availability. Independent of water availability E. globulus maintained a higher A _l :A _s than E. nitens at all ages. Received: 20 March 1997 / Accepted: 30 December 1997     

Sapwood hydraulic conductivity and leaf area – sapwood area relationships following thinning of a Eucalyptus nitens plantation

This study tested the hypothesis that the relationship between leaf area (projected, or one‐sided) and sapwood area in Eucalyptus nitens (Deane and Maiden) Maiden is affected by thinning treatment. However, no difference in the relationship between leaf area and sapwood area was found 8 years after thinning. This result suggests that a single regression equation can be used to predict the leaf area of trees in thinned and unthinned stands. The relationship was non‐linear, implying a causal relationship between growth rate and the ratio of leaf area to sapwood area (A_l : A_s). Sapwood hydraulic conductivity increased by approximately 100% from breast height to crown base, whereas sapwood area decreased by 19%. This implies that the efficiency of water transport through the sapwood increased by 60% along this length. This conclusion is supported by the A_l : A_s relationships which showed that the sapwood area at crown base supported, on average, close to 60% more leaf area than a similar amount of sapwood area at breast height. That large trees in this study had greater hydraulic conductivity and higher A_l : A_s lends support to the argument that resource capture, and hence growth rate, influence sapwood hydraulic conductivity.     

Inter-species variation of photosynthetic and xylem hydraulic traits in the deciduous and evergreen Euphorbiaceae tree species from a seasonally tropical forest in south-western China

The objective of the present study was to examine the functional coordination among hydraulic traits, xylem characteristics and gas exchange rates across three deciduous Euphorbiaceae tree species (Hevea brasiliensis, Macaranga denticulata and Bischofia javanica) and three evergreen Euphorbiaceae tree species (Drypetes indica, Aleurites moluccana and Codiaeum variegatum) from a seasonally tropical forest in south-western China. The deciduous tree species were more vulnerable to water stress-induced embolism than the evergreen tree species. However, the deciduous tree species generally had higher maximal rates of sapwood and leaf-specific hydraulic conductivity (K _S and K _L), respectively. Compared with the evergreen tree species, the deciduous tree species, however, possessed a lower density of sapwood and a wider diameter of xylem vessels. Regardless of leaf phenology, the hydraulic vulnerability and conductivity were significantly correlated with sapwood density and mean vessel diameter. Furthermore, the hydraulic vulnerability was positively correlated with water transport efficiency. In addition, the deciduous tree species exhibited higher maximal photosynthetic rates (A _max) and stomatal conductance (g _max), but lower water use efficiency (WUE). Interestingly, the A _max, g _max and WUE were strongly correlated with K _S and K _L across the deciduous and evergreen tree species. These results suggest that xylem structure, rather than leaf phenology, accounts for the difference in hydraulic traits between the deciduous tree species and the evergreen tree species. Meanwhile, our results show that there is a significant trade-off between hydraulic efficiency and safety, and a strong functional correlation between the hydraulic capacity and gas exchange rates across the deciduous and evergreen tree species.     

Leaf photosynthetic traits scale with hydraulic conductivity and wood density in Panamanian forest canopy trees

We investigated how water transport capacity, wood density and wood anatomy were related to leaf photosynthetic traits in two lowland forests in Panama. Leaf-specific hydraulic conductivity (k_L) of upper branches was positively correlated with maximum rates of net CO₂ assimilation per unit leaf area (A_area) and stomatal conductance (g_s) across 20 species of canopy trees. Maximum k_L showed stronger correlation with A_area than initial k_L suggesting that allocation to photosynthetic potential is proportional to maximum water transport capacity. Terminal branch k_L was negatively correlated with A_area/g_s and positively correlated with photosynthesis per unit N, indicating a trade-off of efficient use of water against efficient use of N in photosynthesis as water transport efficiency varied. Specific hydraulic conductivity calculated from xylem anatomical characteristics (k_theoretical) was positively related to A_area and k_L, consistent with relationships among physiological measurements. Branch wood density was negatively correlated with wood water storage at saturation, k_L, A_area, net CO₂ assimilation per unit leaf mass (A_mass), and minimum leaf water potential measured on covered leaves, suggesting that wood density constrains physiological function to specific operating ranges. Kinetic and static indices of branch water transport capacity thus exhibit considerable co-ordination with allocation to potential carbon gain. Our results indicate that understanding tree hydraulic architecture provides added insights to comparisons of leaf level measurements among species, and links photosynthetic allocation patterns with branch hydraulic processes.     

The effects of intervessel pit characteristics on xylem hydraulic efficiency and photosynthesis in hemiepiphytic and non‐hemiepiphytic Ficus species

Xylem vulnerability to cavitation and hydraulic efficiency are directly linked to fine‐scale bordered pit features in water‐conducting cells of vascular plants. However, it is unclear how pit characteristics influence water transport and carbon economy in tropical species. The primary aim of this study was to evaluate functional implications of changes in pit characteristics for water relations and photosynthetic traits in tropical Ficus species with different growth forms (i.e. hemiepiphytic and non‐hemiepiphytic) grown under common conditions. Intervessel pit characteristics were measured using scanning electron microscopy in five hemiepiphytic and five non‐hemiepiphytic Ficus species to determine whether these traits were related to hydraulics, leaf photosynthesis, stomatal conductance and wood density. Ficus species varied greatly in intervessel pit structure, hydraulic conductivity and leaf physiology, and clear differences were observed between the two growth forms. The area and diameter of pit aperture were negatively correlated with sapwood‐specific hydraulic conductivity, mass‐based net assimilation rate, stomatal conductance (g_s), intercellular CO₂ concentration (C_i) and the petiole vessel lumen diameters (D_v), but positively correlated with wood density. Pit morphology was only negatively correlated with sapwood‐ and leaf‐specific hydraulic conductivity and D_v. Pit density was positively correlated with g_s, C_i and D_v, but negatively with intrinsic leaf water‐use efficiency. Pit and pit aperture shape were not significantly correlated with any of the physiological traits. These findings indicate a significant role of pit characteristics in xylem water transport, carbon assimilation and ecophysiological adaptation of Ficus species in tropical rain forests.     

Coordination of leaf and stem water transport properties in tropical forest trees

Stomatal regulation of transpiration constrains leaf water potential (Ψ_L) within species-specific ranges that presumably avoid excessive tension and embolism in the stem xylem upstream. However, the hydraulic resistance of leaves can be highly variable over short time scales, uncoupling tension in the xylem of leaves from that in the stems to which they are attached. We evaluated a suite of leaf and stem functional traits governing water relations in individuals of 11 lowland tropical forest tree species to determine the manner in which the traits were coordinated with stem xylem vulnerability to embolism. Stomatal regulation of Ψ_L was associated with minimum values of water potential in branches (Ψ_br) whose functional significance was similar across species. Minimum values of Ψ_br coincided with the bulk sapwood tissue osmotic potential at zero turgor derived from pressure–volume curves and with the transition from a linear to exponential increase in xylem embolism with increasing sapwood water deficits. Branch xylem pressure corresponding to 50% loss of hydraulic conductivity (P ₅₀) declined linearly with daily minimum Ψ_br in a manner that caused the difference between Ψ_br and P ₅₀ to increase from 0.4 MPa in the species with the least negative Ψ_br to 1.2 MPa in the species with the most negative Ψ_br. Both branch P ₅₀ and minimum Ψ_br increased linearly with sapwood capacitance (C) such that the difference between Ψ_br and P ₅₀, an estimate of the safety margin for avoiding runaway embolism, decreased with increasing sapwood C. The results implied a trade-off between maximizing water transport and minimizing the risk of xylem embolism, suggesting a prominent role for the buffering effect of C in preserving the integrity of xylem water transport. At the whole-tree level, discharge and recharge of internal C appeared to generate variations in apparent leaf-specific conductance to which stomata respond dynamically.