Developmental patterns of above-ground hydraulic conductance in a Scots pine (Pinus sylvestris L.) age sequence

Hydraulic resistance to water flow was measured in branches and stems of Scots pine trees ranging from 7 to 59 years of age in Thetford (East Anglia, UK). On the basis of these measurements, tree above-ground conductance was calculated and related to the amount of leaf area sustained by each tree. Branches at the crown bottom had a lower proportion of sapwood area and a lower total hydraulic conductance than branches of the same diameter at the tree top. Within branches, most of the hydraulic resistance was located near the needles. Tree above-ground conductance was positively related to tree diameter and inversely related to tree height. Compared with young trees, mature trees had about 4 times less above-ground conductance per unit of leaf area. Apparently, the increase in pathway length associated with tree height growth could be only partially compensated for by the increase in conductive capacities resulting from diameter growth. We argue that this reduction may account for reported decreases of stomatal conductance with tree age. It is suggested that the increase in branchiness associated with tree maturation may represent a compensation mechanism to reduce the overall resistance to water flow in the crowns.     

The ecological significance of long-distance water transport: short-term regulation, long-term acclimation and the hydraulic costs of stature across plant life forms

Plant hydraulic conductance, namely the rate of water flow inside plants per unit time and unit pressure difference, varies largely from plant to plant and under different environmental conditions. Herein the main factors affecting: (a) the scaling between whole‐plant hydraulic conductance and leaf area; (b) the relationship between gas exchange at the leaf level and leaf‐specific xylem hydraulic conductance; (c) the short‐term physiological regulation of plant hydraulic conductance under conditions of ample soil water, and (d) the long‐term structural acclimation of xylem hydraulic conductance to changes in environmental conditions are reviewed. It is shown that plant hydraulic conductance is a highly plastic character that varies as a result of multiple processes acting at several time scales. Across species ranging from coniferous and broad‐leaved trees to shrubs, crop and herbaceous species, and desert subshrubs, hydraulic conductance scaled linearly with leaf area, as expected from first principles. Despite considerable convergence in the scaling of hydraulic properties, significant differences were apparent across life forms that underlie their different abilities to conduct gas exchange at the leaf level. A simple model of carbon allocation between leaves and support tissues explained the observed patterns and correctly predicted the inverse relationships with plant height. Therefore, stature appears as a fundamental factor affecting gas exchange across plant life forms. Both short‐term physiological regulation and long‐term structural acclimation can change the levels of hydraulic conductance significantly. Based on a meta‐analysis of the existing literature, any change in environmental parameters that increases the availability of resources (either above‐ or below‐ground) results in the long‐term acclimation of a less efficient (per unit leaf area) hydraulic system.     

Size-dependency in hydraulic and photosynthetic properties of three <Emphasis Type="Italic">Acer</Emphasis> species having different maximum sizes

Recent studies suggest that physiological traits can be affected by tree size due to stronger hydraulic limitation in taller trees. As trees vary greatly in size, both within and among species, the adaptive responses to hydraulic limitation may be different among species with different maximum sizes. To investigate this, we explored size-dependency in photosynthetic and hydraulic parameters of three Acer species (Acer mono Maxim., Acer amoenum Carr and Acer japonicum Thunb.) using trees of various sizes under well-lit conditions. Leaf stomatal conductance of the Acer species decreased with tree size, implying that water supply to leaves decreases as trees grow. In contrast, content of nitrogen increased with tree size, which may compensate for the decrease in stomatal conductance to maintain the photosynthetic rate. Although the increase in nitrogen and leaf mass per area were larger in species with larger statures, the size-dependency in stomatal conductance was not different among species, and photosynthetic rate and hydraulic conductance were maintained in the three Acer species. Therefore, we suggest that hydraulic limitation on gas exchange does not necessarily depend on the maximum height of the species and that maintenance of photosynthesis and hydraulic properties is a fundamental physiological process during tree growth.     

Bundle sheath lignification mediates the linkage of leaf hydraulics and venation

The lignification of the leaf vein bundle sheath (BS) has been observed in many species and would reduce conductance from xylem to mesophyll. We hypothesized that lignification of the BS in lower‐order veins would provide benefits for water delivery through the vein hierarchy but that the lignification of higher‐order veins would limit transport capacity from xylem to mesophyll and leaf hydraulic conductance (K_leaf). We further hypothesized that BS lignification would mediate the relationship of K_leaf to vein length per area. We analysed the dependence of K_leaf, and its light response, on the lignification of the BS across vein orders for 11 angiosperm tree species. Eight of 11 species had lignin deposits in the BS of the midrib, and two species additionally only in their secondary veins, and for six species up to their minor veins. Species with lignification of minor veins had a lower hydraulic conductance of xylem and outside‐xylem pathways and lower K_leaf. K_leaf could be strongly predicted by vein length per area and highest lignified vein order (R² = .69). The light‐response of K_leaf was statistically independent of BS lignification. The lignification of the BS is an important determinant of species variation in leaf and thus whole plant water transport.     

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     

Safety and efficiency conflicts in hydraulic architecture: scaling from tissues to trees

Tree hydraulic architecture exhibits patterns that propagate from tissue to tree scales. A challenge is to make sense of these patterns in terms of trade-offs and adaptations. The universal trend for conduits per area to decrease with increasing conduit diameter below the theoretical packing limit may reflect the compromise between maximizing the area for conduction versus mechanical support and storage. Variation in conduit diameter may have two complementary influences: one being compromises between efficiency and safety and the other being that conduit tapering within a tree maximizes conductance per growth investment. Area-preserving branching may be a mechanical constraint, preventing otherwise more efficient top-heavy trees. In combination, these trends beget another: trees have more, narrower conduits moving from trunks to terminal branches. This pattern: (1) increases the efficiency of tree water conduction; (2) minimizes (but does not eliminate) any hydraulic limitation on the productivity or tissue growth with tree height; and (3) is consistent with the scaling of tree conductance and sap flow with size. We find no hydraulic reason why tree height should scale with a basal diameter to the two-thirds power as recently claimed; it is probably another mechanical constraint as originally proposed. The buffering effect of capacitance on the magnitude of transpiration-induced xylem tension appears to be coupled to cavitation resistance, possibly alleviating safety versus efficiency trade-offs.     

Growth dynamics of root and shoot hydraulic conductance in seedlings of five neotropical tree species: scaling to show possible adaptation to differing light regimes

The dynamics of growth (shoot and root dry weights, surface areas, hydraulic conductances, and root length) were measured in seedlings of five neotropical tree species aged 4–16 months. The species studied included two light-demanding pioneers (Miconia argentea and Apeiba membranacea) and three shade-tolerant young- or old-forest species (Pouteria reticulata, Gustavia superba, and Trichilia tuberculata). Growth analysis revealed that shoot and root dry weights and hydraulic conductances and leaf area all increased exponentially with time. Alternative methods of scaling measured parameters to reveal differences that might explain adaptations to microsites are discussed. Scaling root conductance to root surface area or root length revealed a few species differences but nothing that correlated with adaptation to light regimes. Scaling of root surface area or root length to root dry weight revealed that pioneers produced significantly more root area and length per gram dry weight investment than shade-tolerant species. Scaling of root and shoot hydraulic conductances to leaf area and scaling of root conductance to root dry weight and shoot conductance to shoot dry weight also revealed that pioneers were significantly more conductive to water than shade-tolerant species. The advantages of scaling hydraulic parameters to leaf surface area are discussed in terms of the Ohm's law analogue of water flow in plants. Received: 24 March 1997 / Accepted: 17 November 1997     

Turning over a new ‘leaf’: multiple functional significances of leaves versus phyllodes in Hawaiian Acacia koa

Hawaiian endemic tree Acacia koa is a model for heteroblasty with bipinnately compound leaves and phyllodes. Previous studies suggested three hypotheses for their functional differentiation: an advantage of leaves for early growth or shade tolerance, and an advantage of phyllodes for drought tolerance. We tested the ability of these hypotheses to explain differences between leaf types for potted plants in 104 physiological and morphological traits, including gas exchange, structure and composition, hydraulic conductance, and responses to varying light, intercellular CO₂, vapour pressure deficit (VPD) and drought. Leaf types were similar in numerous traits including stomatal pore area per leaf area, leaf area‐based gas exchange rates and cuticular conductance. Each hypothesis was directly supported by key differences in function. Leaves had higher mass‐based gas exchange rates, while the water storage tissue in phyllodes contributed to greater capacitance per area; phyllodes also showed stronger stomatal closure at high VPD, and higher maximum hydraulic conductance per area, with stronger decline during desiccation and recovery with rehydration. While no single hypothesis completely explained the differences between leaf types, together the three hypotheses explained 91% of differences. These findings indicate that the heteroblasty confers multiple benefits, realized across different developmental stages and environmental contexts.     

The effect of tree height on crown level stomatal conductance

Variation in stomatal conductance is typically explained in relation to environmental conditions. However, tree height may also contribute to the variability in mean stomatal conductance. Mean canopy stomatal conductance of individual tree crowns (G_Si) was estimated using sap flux measurements in Fagus sylvatica L., and the hypothesis that G_Si decreases with tree height was tested. Over 13 d of the growing season during which soil moisture was not limiting, G_Si decreased linearly with the natural logarithm of vapour pressure deficit (D), and increased exponentially to saturation with photosynthetic photon flux density (Q_o). Under conditions of D = 1 kPa and saturating Q_o, G_Si decreased by approximately 60% with 30 m increase in tree height. Over the same range in height, sapwood‐to‐leaf area ratio (A_S:A_L) doubled. A simple hydraulic model explained the variation in G_Si based on an inverse relationship with height, and a linear relationship with A_S:A_L. Thus, in F. sylvatica, adjustments in A_S:A_L partially compensate for the negative effect of increased flow‐path length on leaf conductance. Furthermore, because stomata with low conductance are less sensitive to D, gas exchange of tall trees is reduced less by high D. Despite these compensations, decreasing hydraulic conductance with tree height in F. sylvatica reduces carbon uptake through a corresponding decrease in stomatal conductance.     

Estimation of hydraulic conductance within field-grown apricot using sap flow measurements

Using the heat pulse and other techniques, the hydraulic architecture of apricot trees was mapped out. The flows (overall flow, flow across the four main branches) and forces (water potential differences between xylem and leaves) measured allowed us to quantify hydraulic conductance of branches and of the root/soil resistance. The experiment was carried out in a commercial orchard of 11-year-old apricot trees (Prunus armeniaca L., cv. Búlida, on Real Fino apricot rootstock) during 1 week (October 27–November 3, 1998). Three representative trees with a cylindrical trunk divided into four main branches of different sizes, orientation and local microclimate were chosen for the experiment. Sap flow was measured throughout the experimental period. Twelve sets of heat-pulse probes were used, one for each main branch. The diurnal course of the environmental conditions, the fraction of the area irradiated and leaf water relations were also considered in each main branch. The relationships between leaf water potential, xylem water potential and transpiration were established for different branches and also for the total plant. Using the slopes of these regressions, total plant conductance, the hydraulic conductance of the stem and root pathway, the hydraulic conductance of the canopy and the hydraulic conductance of each branch were estimated. Our findings show that the root conductance and the canopy hydraulic conductance are similar in magnitude. Leaf hydraulic conductance per leaf area unit was similar for each of the four branch orientations, indicating that, while the light microclimate has a dominant influence on transpiration, in this case it had little effect on the hydraulic properties of the canopy.     

陕北沙地高龄小叶杨光合速率下降的水力限制

黄土高原地区“小老树”现象多出现在成年树,幼树相对较少.为探讨树龄影响“小老树”形成的机制,以该地区“小老树”发生面积最大的树种小叶杨为例,研究了3个不同树龄(低龄:13～15 a;中龄:31～34 a;高龄:49～54 a)树木的生长、光合、水力学特性.结果表明: 随树龄增加,小叶杨枯稍长度显著增加,叶片光合速率、气孔导度和蒸腾速率显著下降,整株水力导度也显著下降,不同测定时间的光合速率、气孔导度与整株水力导度呈显著正相关,表明树龄增加引起的光合速率下降可能与整株水力导度下降有关.与中、低龄相比,高龄小叶杨枝干和叶片抵抗空穴化的能力(P₅₀)更强,但通过脆弱性曲线估算的不同树龄正午时的枝干栓塞程度和叶片水力导度无显著差异,表明高龄小叶杨土-根系统水流阻力的增加可能是导致其整株水力导度降低的重要原因.     

A test of the hydraulic limitation hypothesis in fast-growing Eucalyptus saligna

The hydraulic limitation hypothesis proposes that (1) reduced growth in taller trees is caused by decreased photosynthesis resulting from a decrease in hydraulic conductance promoted by a longer root‐to‐leaf flow path, and (2) this mechanism reduces stand productivity after canopy closure. This hypothesis was tested by comparing the physiology of 7 m (1 year) and 26 m (5 year) Eucalyptus saligna plantations where above‐ground productivity for the 26 m trees was approximately 69% of that for the 7 m trees, and water and nutrients were not limiting. The study compared whole tree physiology [water flux (Q_l), average crown conductance (G_T), crown hydraulic conductance per unit leaf area (K_L), carbon isotope discrimination (δ¹³C)] and leaf physiology under light saturation (leaf water potential at the canopy top (Ψ_LEAF), photosynthetic capacity (A_max), and photosynthesis (A) and stomatal conductance (g_s). K_L was 50% lower in the taller trees, but whole tree Q_l and G_T were the same for the 7 m and 26 m trees. Photosynthetic capacity was the same for leaves at the canopy top, but δ¹³C was ?1.8‰ lower for the 26 m trees. A and g_s were either lower in the taller trees or equal, depending on sampling date. The taller trees maintained 0.8 MPa lower Ψ_LEAF during the day and had 2.6‐times higher sapwood area per unit leaf area; these factors compensated for the effects of increased height and gravitational potential in the taller trees to maintain higher G_T. The hydraulic limitation hypothesis (as originally stated) failed to explain the sharp decline in net primary productivity after canopy closure in this study. The effects of increased height appear to be a universal hydraulic problem for trees, but compensation mitigated these effects and maintained Q_l and G_T in the present study. Compensation may induce other problems (such as lower Ψ_LEAF or higher respiratory costs) that could reduce carbon gain or shift carbon allocation, and future studies of hydraulic limitation should consider compensation and associated carbon costs. In this study, the combination of similar G_T and lower δ¹³C for the 26 m trees suggests that total crown photosynthesis was lower for the 26 m trees, perhaps a result of the lower Ψ_LEAF.     

Comparative hydraulic and anatomic properties in palm trees (Washingtonia robusta) of varying heights: implications for hydraulic limitation to increased height growth

As trees grow taller, the energetic cost of moving water to the leaves becomes higher and could begin to limit carbon gain and subsequent growth. The hydraulic limitation hypothesis states that as trees grow taller, the path length and therefore frictional resistance of water flow increases, leading to stomatal closure, reduced photosynthesis and decreased height growth in tall trees. Although this hypothesis is supported by the physical laws governing water movement in trees, its validation has been complicated by the complex structure of most tree species. Therefore, this study tested the hydraulic limitation hypothesis in Washingtonia robusta (H. Wendl.), a palm that, while growing to tall heights, is still structurally simple enough to act as a model organism for testing. There were no discernable relationships between tree height and stomatal conductance, stomatal densities, guard cell lengths, leaf dry mass per unit area (LMA) or sap flux, suggesting that these key aspects of hydraulic limitation are not reduced in taller palms. Taller palms did, however, have higher maximum daily photosynthetic assimilation rates, lower minimum leaf water potentials that occurred earlier in the day and fewer, smaller leaves than did shorter palms. Leaf epidermal cells were also smaller in taller palms compared with shorter ones. These findings are consistent with hydraulic compensation in that tall palms may be overcoming the increased path length resistance through smaller, more efficient leaves and lower leaf water potentials than shorter palms.     

The hydraulic limitation hypothesis revisited

We proposed the hydraulic limitation hypothesis (HLH) as a mechanism to explain universal patterns in tree height, and tree and stand biomass growth: height growth slows down as trees grow taller, maximum height is lower for trees of the same species on resource-poor sites and annual wood production declines after canopy closure for even-aged forests. Our review of 51 studies that measured one or more of the components necessary for testing the hypothesis showed that taller trees differ physiologically from shorter, younger trees. Stomatal conductance to water vapour (g(s)), photosynthesis (A) and leaf-specific hydraulic conductance (K L) are often, but not always, lower in taller trees. Additionally, leaf mass per area is often greater in taller trees, and leaf area:sapwood area ratio changes with tree height. We conclude that hydraulic limitation of gas exchange with increasing tree size is common, but not universal. Where hydraulic limitations to A do occur, no evidence supports the original expectation that hydraulic limitation of carbon assimilation is sufficient to explain observed declines in wood production. Any limit to height or height growth does not appear to be related to the so-called age-related decline in wood production of forests after canopy closure. Future work on this problem should explicitly link leaf or canopy gas exchange with tree and stand growth, and consider a more fundamental assumption: whether tree biomass growth is limited by carbon availability.     

Plant morphology and root hydraulics are altered by nutrient deficiency in Pistacia lentiscus (L.)

The plants in arid and semiarid areas are often limited by water and nutrients. Morpho-functional adjustments to improve nutrient capture may have important implications on plant water balance, and on plant capacity to withstand drought. Several studies have shown that N and P deficiencies may decrease plant hydraulic conductance. Surprisingly, studies on the implications of nutrient limitations on water use in xerophytes are scarce. We have evaluated the effects of strong reductions in nitrogen and phosphorus availability on morphological traits and hydraulic conductance in seedlings of a common Mediterranean shrub, Pistacia lentiscus L.. Nitrogen deficiency resulted in a decrease in aboveground biomass accumulation, but it did not affect belowground biomass accumulation or root morphology. Phosphorus-deficient plants showed a decrease in leaf area, but no changes in aboveground biomass. Root length, root surface area, and specific root length were higher in phosphorus-deficient plants than in control plants. Nitrogen and phosphorus deficiency reduced both root hydraulic conductance and root hydraulic conductance scaled by total root surface area. On the other hand, nutrient limitations did not significantly affect root conductance per unit of foliar surface area. Thus, adaptation to low nutrient availability did not affect seedling capacity for maintaining water supply to leaves. The implications for drought resistance and survival during seedling establishment in semi-arid environments are discussed.     

A comparison of plant hydraulic conductances in wheat and lupins

Previous studies have shown similar water use for lupins (Lupinusangustifolius L.) and wheat (Triticum aestivum L.), despitea considerably smaller root system in lupins. A field studyand an experiment under controlled conditions using pressure-fluxrelationships were conducted to examine whether higher hydraulicconductances were responsible for the greater water uptake perunit root length in lupins. In the field experiment, the fluxof water and differences in water potential through the soil-plantsystem were measured for both species and used to calculatethe hydraulic conductance through the plant and through theroot and shoot. The hydraulic conductance for the whole plantwas 3–5 times greater in lupins than in wheat. This relativedifference between the species was similar when plant hydraulicconductance was expressed per unit of root length. This occurreddespite the difference in midday water potential between soiland leaves, being consistently greater in wheat (–1.0MPa) than in lupins (–0.7 MPa). When the total plant conductancewas separated into its components, the combined soil and rootconductance and the shoot conductance were 2 and 6 times greater,respectively, in lupins than in wheat. In the experiment undercontrolled conditions, hydraulic conductance for the entireroot system was determined using a pressure chamber. The specificroot hydraulic conductances were 4 times greater in lupins thanin wheat. The results from both field and controlled conditionsexperiments suggest that the greater water uptake per unit rootlength in lupins compared to wheat results from appreciablylarger root and shoot hydraulic conductances. Key words: Lupins, wheat, hydraulic conductances, water, uptake, pressure-flux     

Convergence of tree water use within an arid-zone woodland

We examined spatial and temporal patterns of tree water use and aspects of hydraulic architecture in four common tree species of central Australia—Corymbia opaca, Eucalyptus victrix, E. camaldulensis and Acacia aneura—to better understand processes that constrain water use in these environments. These four widely distributed species occupy contrasting niches within arid environments including woodlands, floodplains and riparian environments. Measurements of tree water use and leaf water potential were made at two sites with contrasting water table depths during a period of high soil water availability following summer rainfall and during a period of low soil water availability following 7 months of very little rainfall during 2007. There were significant differences in specific leaf area (SLA), sapwood area to leaf area ratios and sapwood density between species. Sapwood to leaf area ratio increased in all species from April to November indicating a decline in leaf area per unit sapwood area. Despite very little rainfall in the intervening period three species, C. opaca, E. victrix and E. camaldulensis maintained high leaf water potentials and tree water use during both periods. In contrast, leaf water potential and water use in the A. aneura were significantly reduced in November compared to April. Despite contrasting morphology and water use strategies, we observed considerable convergence in water use among the four species. Wood density in particular was strongly related to SLA, sapwood area to leaf area ratios and soil to leaf conductance, with all four species converging on a common relationship. Identifying convergence in hydraulic traits can potentially provide powerful tools for scaling physiological processes in natural ecosystems.     

豆科复叶和单叶树种的光合-水分关系分析

以豆科（Fabaceae）11个复叶树种和6个单叶树种为材料,测定他们的气孔导度、叶片水力导度、水势、相对含水量等指标,分析叶型对枝叶光合水分关系的影响。结果显示,复叶树种正午叶轴水势（-0.91 MPa）与单叶树种正午枝条水势（-0.88 MPa）间无显著差异,但正午枝条水势（-0.60 MPa）显著高于单叶树种。复叶树种正午气孔导度降低的百分比（55.3%）显著高于单叶树种（34.1%）。叶片、叶轴和枝条正午水势两两之间均显著正相关,但与正午气孔导度之间均不存在相关性。本研究中,17个树种的正午叶片水力导度与气孔导度间显著正相关（r=0.79,P<0.001）,但他们与气孔导度降低百分比间呈负相关（r=-0.81,P<0.001）,说明叶片导水率对日间气孔导度的维持具有重要作用。研究结果表明单叶和复叶树种在光合水分关系上存在明显差异,说明他们对环境条件具有不同的适应策略。     

Size-dependent mortality in a Neotropical savanna tree: the role of height-related adjustments in hydraulic architecture and carbon allocation

Size-related changes in hydraulic architecture, carbon allocation and gas exchange of Sclerolobium paniculatum (Leguminosae), a dominant tree species in Neotropical savannas of central Brazil (Cerrado), were investigated to assess their potential role in the dieback of tall individuals. Trees greater than ∼6-m-tall exhibited more branch damage, larger numbers of dead individuals, higher wood density, greater leaf mass per area, lower leaf area to sapwood area ratio (LA/SA), lower stomatal conductance and lower net CO₂ assimilation than small trees. Stem-specific hydraulic conductivity decreased, while leaf-specific hydraulic conductivity remained nearly constant, with increasing tree size because of lower LA/SA in larger trees. Leaves were substantially more vulnerable to embolism than stems. Large trees had lower maximum leaf hydraulic conductance ( K _leaf) than small trees and all tree sizes exhibited lower K _leaf at midday than at dawn. These size-related adjustments in hydraulic architecture and carbon allocation apparently incurred a large physiological cost: large trees received a lower return in carbon gain from their investment in stem and leaf biomass compared with small trees. Additionally, large trees may experience more severe water deficits in dry years due to lower capacity for buffering the effects of hydraulic path-length and soil water deficits.     

Hydraulic architecture of leaf blades: where is the main resistance?

The hydraulic architecture of Laurus nobilis L. and Juglans regia L. leaves was studied using three different approaches: (1) hydraulic measurements of both intact leaves and of leaves subjected to treatments aimed at removing the extra‐vascular resistance; (2) direct measurements of the vascular pressure with a pressure probe; and (3) modelling the hydraulic architecture of leaf venation system on the basis of measurements of vein densities and conductivities. The hydraulic resistance of leaves (R_leaf) either cut, boiled or frozen–thawed was reduced by about 60 and 85% with respect to control leaves for laurel and walnut, respectively. Direct pressure drop measurements suggested that 88% of the resistance resided outside the vascular system in walnut. Model simulations were in agreement with these results provided vein hydraulic conductance was 0.12–0.28 that of the conductance predicted by Poiseuille's law. The results suggest that R_leaf is dominated by substantial extra‐vascular resistances and therefore contrast with the conclusions of recent studies dealing with the hydraulic architecture of the leaf. The present study confirms the ‘classical’ view of the hydraulic architecture of leaves as composed by a low‐resistance component (the venation) and a high‐resistance component (the mesophyll).