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.     

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。     

Influence of climate-driven shifts in biomass allocation on water transport and storage in ponderosa pine

Conifers decrease the amount of biomass apportioned to leaves relative to sapwood in response to increasing atmospheric evaporative demand. We determined how these climate-driven shifts in allocation affect the aboveground water relations of ponderosa pine growing in contrasting arid (desert) and humid (montane) climates. To support higher transpiration rates, a low leaf:sapwood area ratio (A_L/A_S) in desert versus montane trees could increase leaf-specific hydraulic conductance (K_L). Alternatively, a high sapwood volume:leaf area ratio in the desert environment may increase the contribution of stored water to transpiration. Transpiration and hydraulic conductance were determined by measuring sap flow (J_S) and shoot water potential during the summer (June-July) and fall (August-September). The daily contribution of stored water to transpiration was determined using the lag between the beginning of transpiration from the crown at sunrise and J_S. In the summer, mean maximum J_S was 31.80LJ.74 and 24.34Dž.05 g m^-2 s^-1 for desert and montane trees (a 30.6% difference), respectively. In the fall, J_S was 25.33NJ.52 and 16.36dž.64 g m^-2 s^-1 in desert and montane trees (a 54.8% difference), respectively. J_S was significantly higher in desert relative to montane trees during summer and fall (P<0.05). Predawn and midday shoot water potential and sapwood relative water content did not differ between environments. Desert trees had a 129% higher K_L than montane trees in the summer (2.41᎒^-5 versus 1.05᎒^-5 kg m^-2 s^-1 MPa^-1, P<0.001) and a 162% higher K_L in the fall (1.97᎒^-5 versus 0.75᎒^-5 kg m^-2 s^-1 MPa^-1, P<0.001). Canopy conductance decreased with D in all trees at all measurement periods (P<0.05). Maximum g_C was 3.91 times higher in desert relative to montane trees averaged over the summer and fall. Water storage capacity accounted for 11 kg (11%) and 10.6 kg (17%) of daily transpiration in the summer and fall, respectively, and did not differ between desert and montane trees. By preventing xylem tensions from reaching levels that cause xylem cavitation, high K_L in desert ponderosa pine may facilitate its avoidance. Thus, the primary benefit of low leaf:sapwood allocation in progressively arid environments is to increase K_L and not to increase the contribution of stored water to transpiration.     

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.     

Regulation of xylem sap flow in an evergreen, a semi-deciduous, and a deciduous Meliaceae species from the Amazon

The significance of phenological characteristics, stomatal conductance of the leaves, and stem water storage fluctuations for the regulation of xylem sap flow in an evergreen (Carapa guianensis Aubl.), in a semi-deciduous (Swietenia macrophylla King), and in a deciduous (Cedrela odorata L.) Meliaceae species was studied in a 7-year-old plantation near Manaus, Brazil. The study responds to the increasing demand for knowledge on the water relations of highly exploited timber trees of the Amazon. Xylem sap flow measurements indicated that the daily sap flow of Carapa (3.8 l day^-1 tree^-1 to 16.4 l day^-1 tree^-1) exceeded the daily sap flow of Swietenia (2.4 l day^-1 tree^-1 to 7.0 l day^-1 tree^-1) and Cedrela (1.6 l day^-1 tree^-1 to 11.6 l day^-1 tree^-1) during the entire year, although the highest flux densities were measured in Cedrela. The decrease in xylem sap flow observed in periods with low soil water potentials and high atmospheric vapor saturation deficits was more pronounced in the deciduous (Cedrela) and semi-deciduous species (Swietenia) than the evergreen species (Carapa). Carapa, which has the highest daily sap flow, had the highest biomass and sapwood portion. The high flux densities measured in Cedrela most likely result from the large earlywood vessels in this species. The seasonal variation of xylem sap flow of the three species was correlated with the stomatal conductance of the leaves measured by infiltration experiments. Stem water storage fluctuations in Carapa and Swietenia were predominantly due to transpiration; in Cedrela it was predominantly due to evaporative water loss on the stem surface during dry periods.     

Relations between efficiency of water transport and duration of leaf growth in some deciduous and evergreen trees

Shoot and leaf growth rate as well as shoot hydraulic conductance per unit leaf area (K_SL) were measured on three evergreen (Viburnum tinus L., Prunus laurocerasus L., Laurus nobilis L.) and three deciduous (Corylus avellana L., Juglans regia L., Castanea sativa L.) trees growing under the same environmental conditions. The times required to complete shoot growth (27 days for P. laurocerasus to 51 days for V. tinus) and leaf expansion (24 days for C. sativa to 42 days for C. avellana) were very different among the studied species. These species also differed in K_SL that ranged between 1.5 and 3.5 e-4 kg s^-1 m^-2 MPa^-1 in C. avellana and C. sativa, respectively, with intermediate values recorded in the other species. A strong, negative and statistically significant correlation was found to exist between K_SL and the time required for complete leaf expansion. This suggests that duration of leaf growth is shortened by the high hydraulic efficiency of the shoot. In contrast, no statistically significant relationship was found to exist between K_SL and shoot growth rate. Whether a high leaf growth rate can be interpreted as advantageous to plants or it is only an epiphenomenon of the high efficiency in the vertical water transport is discussed.     

Limitation of stomatal conductance by hydraulic traits: sensing or preventing xylem cavitation?

We tested the hypothesis that hydraulic conductance per unit leaf surface area of plant shoots (K_SL) determines the maximum diurnal stomatal conductance (g_L) that can be reached by plants growing in the field. A second hypothesis was tested that some xylem cavitation cannot be avoided by transpiring plants and might act as a signal for regulating g_L. Eleven woody species were studied, differing from each other with respect to taxonomy, wood anatomy and leaf habit. Maximum diurnal g_L, transpiration rate (E_L), pre-dawn and minimum diurnal leaf water potential (O_pd and O_min, respectively) were measured in the field. The critical O level at which stem cavitation was triggered (O_cav) was measured on detached branches, using the acoustic method. A high-pressure flow meter was used to measure maximum K_SL of 1-year-old shoots. Both g_L and E_L were positively related to K_SL. The whole-plant hydraulic conductance per unit leaf area (K_WL) of all the species studied, calculated as the ratio of E_L to (O (=O_pd-O_min) was closely related to K_SL. In every case, O_min (ranging between -0.85 and -1.35 MPa in the different species) dropped to the O_cav range or was <O_cav (ranging between -0.71 and -1.23 MPa), thus suggesting that some cavitation-induced embolism could not be avoided. The possibility is discussed that some cavitation-induced reduction in K_SL is the signal for stomatal closure preventing runaway embolism. The lack of correlation of g_L to O_cav is discussed in terms of the inconsistency of O_cav as an indicator of the vulnerability of plants to cavitation. No differences in hydraulic traits were observed between evergreen and deciduous species.     

Responses of sap flux and stomatal conductance of Pinus taeda L. trees to stepwise reductions in leaf area

Herbivory or artificial foliage removal has been shown to affect gas exchange and canopy water relations. In this study, canopy architecture and water relations in response to progressive defoliation were examined in a stand of 8-year-old loblolly pine (Pinus taeda L.) trees, a shade-intolerant, pioneer species common in the south-eastern USA. Sap flux was measured with constant heat sap flow gauges in order to estimate canopy stomatal conductance (Gs) while foliage in the 6 m high stand was harvested in 1 m increments from the bottom up. Leaf-level stomatal conductance and water potential data were also collected. Profiles of silhouette area ratio and specific leaf area showed no trends with crown height, reflecting an open canopy (leaf area index = 1.55). Therefore, short-term changes in Gs with foliage removal were attributed to hydraulic effects rather than influences of changes in mean microclimate conditions on Gs of remaining foliage. A large increase in Gs was observed during the 6 h pruning period which fully compensated for the reductions in foliage area down to 45%. Canopy stomatal conductance and whole plant liquid phase conductance as calculated from sap flux were both influenced by the rate of growth as indicated by the annual basal area increment.     

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.     

Does foliage on the same branch compete for the same water? Experiments on Douglas-fir trees

Do branchlets within a branch have autonomous water supplies, or do they share a common water supply system? We hypothesized that if branchlets shared a common water supply, then stomatal conductance (g_s) on sunlit foliage would increase with reduced transpiration of competing foliage on the branch. We reduced transpiration of other foliage on the branch through bagging and shading, and we monitored the gas-exchange responses of the remaining sunlit foliage on the branch relative to control branches for several age classes of Douglas-fir trees (aged ~10 years, 20 years, and 450 years old). Contrary to our hypothesis, we found no increases in g_s in either young or old trees following transient reductions in the amount of transpiring leaf area. The diurnal change in water potential, mid-day stomatal closure and associated photosynthetic decline occurred at the same time and were of the same magnitude on both treated and untreated branches, with the exception of photosynthesis in one 450-year-old tree. Hydraulic conductance measurements of branch junctions indicate that xylem within branches is only partially interconnected which would reduce the effectiveness of shading as a means of increasing water supply to the remaining sunlit foliage. The lack of a response implies that when a branch is in partial shade, the remaining sunlit foliage has no advantage with respect to water status over foliage on a branch completely in the sun.     

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     

Physiological basis of spacing effects on tree growth and form in Eucalyptus globulus

We examine the effects of spacing and layout on the growth and form of 3- to 4-year-old Eucalyptus globulus in a farm forestry context. Four planting layouts were chosen. These represented the range commonly in use in farm forestry: block plantings (2Ǹ m), triple rows (2Ǹ m) at 10-m intervals, single rows (2᎒ m) and isolated trees (10᎒ m). The physiological significance of key results is interpreted in terms of changes in the parameters of a simple plantation growth model. Under conditions where levels of direct light are high, for example during summer, block-planted trees intercepted only 38% of the light intercepted by isolated trees. On a stand basis, however, the combination of incident radiation and ground coverage declined with lower stand densities. While stand leaf area index declined from around 6 to 1 with increased spacing, individual tree leaf areas rose from around 50 m² in block plantings to 150 m² in isolated trees. The proportion of above-ground biomass found in stems declined with increasing spacing as the mass in foliage and branches increased. Stems accounted for 65% of above-ground biomass in block-planted trees but only 35% in isolated trees. The contributions of leaves and branches correspondingly rose from 19% to 35% and from 16% to 29%, respectively. Changes in biomass distribution were accompanied by increasing branch number, branch thickness, flatter branch angles and the longer retention of lower branches with greater spacing. These changes have implications for the merchantability of the timber. The efficiency of above-ground radiation conversion was constant at 0.67 g MJ^-1 irrespective of spacing. We estimated that foliar maintenance respiration (R_m) accounted for about 90% of above-ground R_m. On a stand basis R_m costs block plantings 23.90 t DM ha^-1 year^-1 (50% annual above-ground photosynthetic production) compared with 6.22 t DM ha^-1 year^-1 (40% annual above-ground photosynthetic production) in stands of isolated trees.     

树高对马占相思整树水分利用的效应

利用Granier热消散探针,于2004年观测了华南丘陵坡地常见绿化先锋树种马占相思(22年生)的树干液流,同时监测林冠上方的光合有效辐射、气温、相对湿度和0～30 cm的土壤体积含水量.结合树木的形态特征、液流密度和简化的Whitehead & Jarvis公式,分别计算了整树蒸腾、冠层气孔导度和叶面积/边材面积比值,分析了树高对整树蒸腾、冠层气孔导度和叶面积/边材面积比值的影响.结果表明：土壤水分充足时,马占相思整树蒸腾随树高呈二次多项式增加(P<0.01),冠层气孔导度日变化均呈“单峰”格型;在所有光合有效辐射范围内,高树的参比冠层气孔导度和冠层气孔导度对水汽压亏缺的敏感性均高于矮树;叶面积/边材面积比值为（1.837±0.048） m²·cm^-2,并与树高呈幂函数关系.随着树木高度的增加,马占相思没有发生明显的水力限制和补偿.     

Effect of drought on leaf gland secretion of the mangrove Avicennia germinans L.

This study assessed the effects of salinity changes over space and time upon leaf gland secretion in Avicennia germinans trees growing naturally in an area featuring markedly seasonal rainfall. Soil ? (, soil N MPa) during the wet season was -0.95ǂ.05 and -2.12ǂ.08 at low and high salinity sites, respectively. During the dry season, these values decreased to -3.24ǂ.09 at low salinity and to -5.75ǂ.06 at high salinity. Consequently, predawn and midday plant water potential were lowered during drought at both sites. The rates of secretion (mmol m^-2 h^-1 ) increased during drought from 0.91ǂ.12 during the wet season to 1.93ǂ.12 at low salinity, and from 1.69ǂ.12 during the wet season to 2.81ǂ.15 at high salinity. Conversely, stomatal conductance (g_s) was lowered by both salinity and drought. As xylem osmolality increased during drought, secretion tended to rise exponentially, and g_s decayed hyperbolically. Thus, a trade-off is obtained between enhancement in salt secretion and control of water loss suggested by g_s.     

Photosynthetic characteristics in canopies of Quercus rubra, Quercus prinus and Acer rubrum differ in response to soil water availability

Photosynthesis and related leaf characteristics were measured in canopies of co-occurring Quercus rubra L. (red oak), Quercus prinus L. (chestnut oak) and Acer rubrum L. (red maple) trees. Mature (20+ m tall) trees were investigated at sites of differing soil water availability within a catchment (a drier upper site and a wetter lower site). Leaf photosynthetic characteristics differed significantly between species and in response to site and position in the canopy. Photosynthetic capacity (A_max) was significantly greater at the wetter site in all canopy strata in A. rubrum but not in Q. rubra or Q. prinus. Our findings for A. rubrum are generally consistent with those predicting that species with higher specific leaf area (SLA) will have higher A_max per unit leaf nitrogen (N) and that species with leaves with lower SLA (e.g. Q. rubra and Q. prinus) will have shallower slopes of the A_max-N relationship. Importantly, the relationships between A_max and N_area (and by implication photosynthetic nitrogen-use efficiency, PNUE) differed in A. rubrum between the sites, with PNUE significantly lower at the drier site. The lower photosynthetic capacity and PNUE must substantially reduce carbon acquisition capacity in A. rubrum under these field conditions. Maximum stomatal conductance (g_smax) differed significantly between species, with g_smax greatest in Q. rubra and Q. prinus. In Q. rubra and Q. prinus, g_smax was significantly lower at the upper site than the lower site. There was no significant response of g_smax to site in A. rubrum. These stomatal responses were consistent with the C_i/C_a ratio, which was significantly lower in leaves of Q. rubra and Q. prinus at the upper site, but did not differ between sites in A. rubrum. Leaf '¹³C was significantly lower in A. rubrum than in either Q. rubra or Q. prinus at both sites. These findings indicate differences in stomatal behaviour in A. rubrum which are likely to contribute to lower water use efficiency at both sites. Our results support the hypothesis that the two Quercus species, in contrast to A. rubrum, maintain photosynthetic capacity at the drier site whilst minimising transpirational water loss. They also suggest, based primarily on physiological evidence, that the ability of A. rubrum to compete with other species of these deciduous forests may be limited, particularly in sites of low moisture availability and during low rainfall years.     

Effects of tree height on branch hydraulics, leaf structure and gas exchange in California redwoods

We examined changes in branch hydraulic, leaf structure and gas exchange properties in coast redwood ( Sequoia sempervirens ) and giant sequoia ( Sequoiadendron giganteum ) trees of different sizes. Leaf-specific hydraulic conductivity ( k _L) increased with height in S. sempervirens but not in S. giganteum , while xylem cavitation resistance increased with height in both species. Despite hydraulic adjustments, leaf mass per unit area (LMA) and leaf carbon isotope ratios ( δ ¹³C) increased, and maximum mass-based stomatal conductance ( g _mass) and photosynthesis ( A _mass) decreased with height in both species. As a result, both A _mass and g _mass were negatively correlated with branch hydraulic properties in S. sempervirens and uncorrelated in S. giganteum . In addition, A _mass and g _mass were negatively correlated with LMA in both species, which we attributed to the effects of decreasing leaf internal CO₂ conductance ( g _i). Species-level differences in wood density, LMA and area-based gas exchange capacity constrained other structural and physiological properties, with S. sempervirens exhibiting increased branch water transport efficiency and S. giganteum exhibiting increased leaf-level water-use efficiency with increasing height. Our results reveal different adaptive strategies for the two redwoods that help them compensate for constraints associated with growing taller, and reflect contrasting environmental conditions each species faces in its native habitat.     

Crown conductance in dwarf, medium, and tall pitch pines in the Long Island Pine Plains

The New York Pine Plains are a unique ecosystem with normal statured and a dwarfed variety of pitch pines (Pinus rigida Mill.). Growing interspersed with the dwarf pines are trees of intermediate height and features. Several hypotheses have been put forward as to why some of the trees are dwarfed, but none have been substantiated. In this study, we tested whether dwarf or medium trees are hydraulically limited compared to normally growing trees. Granier style sap flux sensors were installed in three to six trees of each tree type and sap flux was measured in early August 2004. Sap flux measurements were scaled to crown stomatal conductance using leaf area to sapwood area ratios for each tree. Contrary to expectation, dwarf and medium stature trees had very low leaf area to sapwood area ratios, but high crown stomatal conductances compared to normal trees. Analyses of leaf area, ring widths, and crown stomatal conductance indicate that differences between normal, and dwarf and medium pines are not due to hydraulic limitation, but that stunted growth may be due to other causes.     

Wood anatomy constrains stomatal responses to atmospheric vapor pressure deficit in irrigated, urban trees

Plant transpiration is strongly constrained by hydraulic architecture, which determines the critical threshold for cavitation. Because species vary greatly in vulnerability to cavitation, hydraulic limits to transpiration and stomatal conductance have not generally been incorporated into ecological and climate models. We measured sap flow, leaf transpiration, and vulnerability to cavitation of a variety of tree species in a well-irrigated but semi-arid urban environment in order to evaluate the generality of stomatal responses to high atmospheric vapor pressure deficit (D). We found evidence of broad patterns of stomatal responses to humidity based on systematic differences in vulnerability to cavitation. Ring-porous taxa consistently had vulnerable xylem and showed strong regulation of transpiration in response to D, while diffuse-porous taxa were less vulnerable and transpiration increased nearly linearly with D. These results correspond well to patterns in the distribution of the taxa, such as the prevalence of diffuse-porous species in riparian ecosystems, and also provide a means of representing maximum transpiration rates at varying D in broad categories of trees.     

Water relations of coastal and estuarine Rhizophora mangle: xylem pressure potential and dynamics of embolism formation and repair

Physiological traits related to water transport were studied in Rhizophora mangle (red mangrove) growing in coastal and estuarine sites in Hawaii. The magnitude of xylem pressure potential (P_x), the vulnerability of xylem to cavitation, the frequency of embolized vessels in situ, and the capacity of R. mangle to repair embolized vessels were evaluated with conventional and recently developed techniques. The osmotic potential of the interstitial soil water (?_sw) surrounding the roots of R. mangle was c. -2.6LJ.52᎒^-3 and -0.4Lj.13᎒^-3 MPa in the coastal and estuarine sites, respectively. Midday covered (non-transpiring) leaf water potentials (O_L) determined with a pressure chamber were 0.6-0.8 MPa more positive than those of exposed, freely-transpiring leaves, and osmotic potential of the xylem sap (?_x) ranged from -0.1 to -0.3 MPa. Consequently, estimated midday values of P_x (calculated by subtracting ?_x from covered O_L) were about 1 MPa more positive than O_L determined on freely transpiring leaves. The differences in O_L between covered and transpiring leaves were linearly related to the transpiration rates. The slope of this relationship was steeper for the coastal site, suggesting that the hydraulic resistance was larger in leaves of coastal R. mangle plants. This was confirmed by both hydraulic conductivity measurements on stem segments and high-pressure flowmeter studies made on excised leafy twigs. Based on two independent criteria, loss of hydraulic conductivity and proportions of gas- and liquid-filled vessels in cryo-scanning electron microscope (cryo-SEM) images, the xylem of R. mangle plants growing at the estuarine site was found to be more vulnerable to cavitation than that of plants growing at the coastal site. However, the cryo-SEM analyses suggested that cavitation occurred more readily in intact plants than in excised branches that were air-dried in the laboratory. Cryo-SEM analyses also revealed that, in both sites, the proportion of gas-filled vessels was 20-30% greater at midday than at dawn or during the late afternoon. Refilling of cavitated vessels thus occurred during the late afternoon when considerable tension was present in neighboring vessels. These results and results from pressure-volume relationships suggest that R. mangle adjusts hydraulic properties of the water-transport system, as well as the leaf osmotic potential, in concert with the environmental growing conditions.     

Integrated responses of hydraulic architecture, water and carbon relations of western hemlock to dwarf mistletoe infection

Dwarf mistletoe (Arceuthobium spp.) is a hemiparasite that is said to be the single‐most destructive pathogen of commercially valuable coniferous trees in many regions of the world. Although its destructive nature is well documented in many respects, its effects on the physiology of its host are poorly understood. In the present study, water and carbon relations were characterized over a range of scale from leaf to whole tree in large (40‐ to 50‐m‐tall) individuals of western hemlock (Tsuga heterophylla (Raf.) Sarg.) that were either heavily infected, or uninfected with hemlock dwarf mistletoe (Arceuthobium tsugense). Specific hydraulic conductivity (k_s) of infected branches was approximately half that of uninfected branches, yet leaf‐specific conductivity (k_L) was similar because leaf area : sapwood area ratios (A_L : A_S) of infected branches were lower. Pre‐dawn and minimum leaf water potential and stomatal conductance (g_s) were similar among infected and uninfected trees because adjustments in hydraulic architecture of infected trees maintained k_L despite reduced k_s. Maximum whole‐tree water use was substantially lower in infected trees (approximately 55 kg d^?1) than in uninfected trees (approximately 90 kg d^?1) because reduced numbers of live branches in infected trees reduced whole‐tree A_L : A_S in a manner consistent with that observed in infected branches. Maximum photosynthetic rates of heavily infected trees were approximately half those of uninfected trees. Correspondingly, leaf nitrogen content was 35% lower in infected trees. Foliar δ¹³C values were 2.8‰ more negative in infected than in uninfected individuals, consistent with the absence of stomatal adjustment to diminished photosynthetic capacity. Adjustments in hydraulic architecture of infected trees thus contributed to homeostasis of water transport efficiency and transpiration on a leaf area basis, whereas both carbon accumulation and photosynthetic water use efficiency were sharply reduced at both the leaf and whole‐tree scale.