Xylem conductivity and vulnerability to cavitation of ponderosa pine growing in contrasting climates

We examined the effects of increased transpiration demand on xylem hydraulic conductivity and vulnerability to cavitation of mature ponderosa pine (Pinus ponderosa Laws.) by comparing trees growing in contrasting climates. Previous studies determined that trees growing in warm and dry sites (desert) had half the leaf/sapwood area ratio (A(L)/A(S)) and more than twice the transpiration rate of trees growing in cool and moist sites (montane). We predicted that high transpiration rates would be associated with increased specific hydraulic conductivity (K(S)) and increased resistance to xylem cavitation. Desert trees had 19% higher K(S) than montane trees, primarily because of larger tracheid lumen diameters. Predawn water potential and water potential differences between the soil and the shoot were similar for desert and montane trees, suggesting that differences in tracheid anatomy, and therefore K(S), were caused primarily by temperature and evaporative demand, rather than soil drought. Vulnerability to xylem cavitation did not differ between desert and montane populations. A 50% loss in hydraulic conductivity occurred at water potentials between -2.61 and -2.65 MPa, and vulnerability to xylem cavitation did not vary with stem size. Minimum xylem tensions of desert and montane trees did not drop below -2.05 MPa. Foliage turgor loss point did not differ between climate groups and corresponded to mean minimum xylem tensions in the field. In addition to low A(L)/A(S), high K(S) in desert trees may provide a way to increase tree hydraulic conductivity in response to high evaporative demand and prevent xylem tensions from reaching values that cause catastrophic cavitation. In ponderosa pine, the flexible responses of A(L)/A(S) and K(S) to climate may preclude the existence of significant intraspecific variation in the vulnerability of xylem to cavitation.     

The fate of hydraulically redistributed water in a semi-arid zone eucalyptus species

Although hydraulic redistribution has been observed for a range of tree species, including Eucalyptus kochii subsp. borealis (C. Gardner) D. Nicolle, there is limited direct evidence that water taken up by deep roots in moist soil is in fact exuded by shallow roots in dry soil. This paper reports an experiment designed to test this hypothesis. Water enriched with deuterium was added to the groundwater via a slotted tube at 4.5 m depth below 5-year-old E. kochii subsp. borealis trees. Nocturnal sap flow increased markedly immediately after deep irrigation, indicating that the trees were using water from this depth. Two weeks later, samples of surface soil and xylem water were found to contain levels of deuterium up to 30% higher than soils and xylem water from a control plot upslope of the main treatment plot. This is strong evidence that trees used groundwater and that efflux of important amounts of hydraulically redistributed water occurred via the roots of E. kochii subsp. borealis.     

Tolerance of salinized floodplain conditions in a naturally occurring Eucalyptus hybrid related to lowered plant water potential

Rising saline groundwater and reduced flooding frequency are causing dieback of Eucalyptus largiflorens F. Muell. along the Murray River in Australia. A green-leaved variant of E. largiflorens, which is probably a hybrid with a local mallee species (E. gracilis F. Muell.), tolerates saline conditions better than the more common grey-leaved variant. The green variant exhibited more negative water potentials than the grey variant, and comparison with soil water potential profiles indicated that the green variant extracted water from slightly higher up the soil profile where the salt content was lower but the soil was drier. However, the stable isotopes of water (2H and 18O) in the xylem did not differ significantly between paired green and grey trees, suggesting that both variants used the same water source. The green variant may be able to extract water for a longer period from a given point in the soil profile and tolerate a higher salt concentration around its roots than the grey variant. Predawn leaf water potentials of both variants decreased with increasing salinity of groundwater and decreasing depth to the groundwater, probably because the roots were being progressively confined to soil with lower matric potential as groundwater discharge through transpiration progressively salinized soil up the profile. The green variant had a lower assimilation rate and stomatal conductance than the grey variant, although the differences were not statistically significant during most of the year. Discrimination of 13C indicated that the green variant had a higher leaf internal CO2 concentration than the grey variant, indicative of a greater biochemical limitation on photosynthesis, perhaps resulting from the effects of operating at lower water potentials. The green variant had significantly lower stem hydraulic conductivity than the grey variant, probably because of its smaller xylem vessel diameter and higher degree of embolism. The more conservative water use of the green variant and its ability to operate at lower water potential than the grey variant appear to underlie its ability to tolerate conditions of reduced useable water above the saline groundwater. This advantage appears to outweigh the costs of increased xylem embolism and reduced assimilation.     

Variation in water potential, hydraulic characteristics and water source use in montane Douglas-fir and lodgepole pine trees in southwestern Alberta and consequences for seasonal changes in photosynthetic capacity

Tree species response to climate change-induced shifts in the hydrological cycle depends on many physiological traits, particularly variation in water relations characteristics. We evaluated differences in shoot water potential, vulnerability of branches to reductions in hydraulic conductivity, and water source use between Pinus contorta Dougl. ex Loud. var. latifolia Engelm. (lodgepole pine) and Pseudotsuga menziesii (Mirb.) Franco (interior Douglas-fir), and determined the consequences for seasonal changes in photosynthetic capacity. The Douglas-fir site had soil with greater depth, finer texture and higher organic matter content than soil at the lodgepole pine site, all factors that increased the storage of soil moisture. While the measured xylem vulnerability curves were quite similar for the two species, Douglas-fir had lower average midday shoot water potentials than did lodgepole pine. This implied that lodgepole pine exhibited stronger stomatal control of transpiration than Douglas-fir, which helped to reduce the magnitude of the water potential gradient required to access water from drying soil. Stable hydrogen isotope measurements indicated that Douglas-fir increased the use of groundwater during mid-summer when precipitation inputs were low, while lodgepole pine did not. There was a greater reduction of photosynthetic carbon gain in lodgepole pine compared with Douglas-fir when the two tree species were exposed to seasonal declines in soil water content. The contrasting patterns of seasonal variation in photosynthetic capacity observed for the two species were a combined result of differences in soil characteristics at the separate sites and the inherent physiological differences between the species.     

Does growing on a slope affect tree xylem structure and water relations?

Variations in slope, exposure, relief and substrate over a short distance and their influences on plant function are poorly understood. We investigated the influences of soil hydrological characteristics on internal stem structure and hydraulic properties of downy oak (Quercus pubescens Willd.) growing along a hill slope. Increment wood cores were extracted from the base and at breast height (BH) of tree stems. Relative wood water content (W(c)) and wood density (D(w)) were measured in the sapwood. Wood compression strength (delta) in the longitudinal direction was measured with a fractometer. Thin sections were cut from the transversal face of each core and vessel lumen area (V(A)) was measured and xylem theoretical hydraulic conductivity (L(th)) estimated over the sapwood. Topsoil volumetric water content (theta(v)) was determined around trees and the hydrological behavior of the slope was studied through field surveys. Data were used as input to a hydrological model to simulate topsoil water distribution along the slope. Results showed that theta(v) tends to decrease with increasing altitude. Groundwater levels were lower upslope than downslope, and results from the hydrological model confirmed these trends. Mean W(c) at the base of each tree decreased significantly with increasing altitude, whereas at BH, no differences were found along the slope. There was a significant positive relationship between W(c) measured at the tree base and theta(v) along the hill slope, but not for W(c) measured at BH. Values of D(w) and delta measured at both stem positions increased significantly with increasing altitude and decreasing theta(v). Significant negative relationships were found between delta and theta(v) measured at the stem base and at BH. At both stem positions, delta was closely related to D(w) and L(th). Vessel lumen areas at BH and the stem base were significantly regressed with altitude, theta(v), D(w) and delta. Xylem theoretical hydraulic conductivity at both stem positions was negatively related to altitude and soil theta(v), but only L(th) measured at the stem base was negatively regressed with D(w). The results are discussed in the context of how tree position along a hill slope influences water uptake and internal xylem structure.     

Partitioning of soil water among tree species in a Brazilian Cerrado ecosystem

Source water used by woody perennials in a Brazilian savanna (Cerrado) was determined by comparing the stable hydrogen isotope composition (deltaD) of xylem sap and soil water at different depths during two consecutive dry seasons (1995 and 1996). Plant water status and rates of water use were also determined and compared with xylem water deltaD values. Overall, soil water deltaD decreased with increasing depth in the soil profile. Mean deltaD values were -35 per thousand for the upper 170 cm of soil and -55 per thousand between 230 and 400 cm depth at the end of the 1995 dry season. Soil water content increased with depth, from 18% near the surface to about 28% at 400 cm. A similar pattern of decreasing soil water deltaD with increasing depth was observed at the end of the 1996 dry season. Patterns consistent with hydraulic lift were observed in soil profiles sampled in 1995 and 1997. Concurrent analyses of xylem and soil water deltaD values indicated a distinct partitioning of water resources among 10 representative woody species (five deciduous and five evergreen). Among these species, four evergreen and one deciduous species acquired water primarily in the upper soil layers (above 200 cm), whereas three deciduous and one evergreen species tapped deep sources of soil water (below 200 cm). One deciduous species exhibited intermediate behavior. Total daily sap flow was negatively correlated with xylem sap deltaD values indicating that species with higher rates of water use during the dry season tended to rely on deeper soil water sources. Among evergreen species, minimum leaf water potentials were also negatively correlated with xylem water deltaD values, suggesting that access to more readily available water at greater depth permitted maintenance of a more favorable plant water status. No significant relationship between xylem water deltaD and plant size was observed in two evergreen species, suggesting a strong selective pressure for small plants to rapidly develop a deep root system. The degree of variation in soil water partitioning, leaf phenology and leaf longevity was consistent with the high diversity of woody species in the Cerrado.     

Variation in soil water uptake and its effect on plant water status in Juglans regia L. during dry and wet seasons

Temporal and spatial variations in the water status of walnut trees (Juglans regia L.) and the soil in which they were growing were traced by analyzing the differences in hydrogen isotopes during spring and summer in a 7-year-old walnut stand. Walnut root dynamics were measured in both dry and wet seasons. Walnut roots were mainly distributed in the upper soil (0-30 cm depth), with around 60% of the total root mass in upper soil layers and 40% in deep soil layers (30-80 cm depth). The upper soil layers contributed 68% of the total tree water requirement in the wet season, but only 47% in the dry season. In the wet season, total roots, living roots and new roots were all significantly more abundant than in the dry season. There were significant differences in pre-dawn branch percentage loss of hydraulic conductance (PLC), pre-dawn leaf water potential and transpiration between the dry and wet seasons. Water content in the upper soil layers remarkably influenced xylem water stable-hydrogen isotope (δD) values. Furthermore, there were linear relationships between the xylem water δD value and pre-dawn branch PLC, pre-dawn leaf water potential, transpiration rate and photosynthetic rate. In summary, J. regia was compelled to take a larger amount of water from the deep soil layers in the dry season, but this shift could not prevent water stress in the plant. The xylem water δD values could be used as an indicator to investigate the water stress of plants, besides probing profiles of soil water use.     

Influence of soil texture on hydraulic properties and water relations of a dominant warm-desert phreatophyte

We investigated hydraulic constraints on water uptake by velvet mesquite (Prosopis velutina Woot.) at a site with sandy-loam soil and at a site with loamy-clay soil in southeastern Arizona, USA. We predicted that trees on sandy-loam soil have less negative xylem and soil water potentials during drought and a lower resistance to xylem cavitation, and reach E(crit) (the maximum steady-state transpiration rate without hydraulic failure) at higher soil water potentials than trees on loamy-clay soil. However, minimum predawn leaf xylem water potentials measured during the height of summer drought were significantly lower at the sandy-loam site (-3.5 +/- 0.1 MPa; all errors are 95% confidence limits) than at the loamy-clay site (-2.9 +/- 0.1 MPa). Minimum midday xylem water potentials also were lower at the sandy-loam site (-4.5 +/- 0.1 MPa) than at the loamy-clay site (-4.0 +/- 0.1 MPa). Despite the differences in leaf water potentials, there were no significant differences in either root or stem xylem embolism, mean cavitation pressure or Psi(95) (xylem water potential causing 95% cavitation) between trees at the two sites. A soil-plant hydraulic model parameterized with the field data predicted that E(crit) approaches zero at a substantially higher bulk soil water potential (Psi(s)) on sandy-loam soil than on loamy-clay soil, because of limiting rhizosphere conductance. The model predicted that transpiration at the sandy-loam site is limited by E(crit) and is tightly coupled to Psi(s) over much of the growing season, suggesting that seasonal transpiration fluxes at the sandy-loam site are strongly linked to intra-annual precipitation pulses. Conversely, the model predicted that trees on loamy-clay soil operate below E(crit) throughout the growing season, suggesting that fluxes on fine-textured soils are closely coupled to inter-annual changes in precipitation. Information on the combined importance of xylem and rhizosphere constraints to leaf water supply across soil texture gradients provides insight into processes controlling plant water balance and larger scale hydrologic processes.     

Regulation of transpirational water loss in Quercus suber trees in a Mediterranean-type ecosystem

Sap flux density in branches, leaf transpiration, stomatal conductance and leaf water potentials were measured in 16-year-old Quercus suber L. trees growing in a plantation in southern Portugal to understand how evergreen Mediterranean trees regulate water loss during summer drought. Leaf specific hydraulic conductance and leaf gas exchange were monitored during the progressive summer drought to establish how changes along the hydraulic pathway influence shoot responses. As soil water became limiting, leaf water potential, stomatal conductance and leaf transpiration declined significantly. Predawn leaf water potential reflected soil water potential measured at 1-m depth in the rhizospheres of most trees. The lowest predawn leaf water potential recorded during this period was -1.8 MPa. Mean maximum stomatal conductance declined from 300 to 50 mmol m(-2) s(-1), reducing transpiration from 6 to 2 mmol m(-2) s(-1). Changes in leaf gas exchange were attributed to reduced soil water availability, increased resistances along the hydraulic pathway and, hence, reduced leaf water supply. There was a strong coupling between changes in soil water content and stomatal conductance as well as between stomatal conductance and leaf specific hydraulic conductance. Despite significant seasonal differences among trees in predawn leaf water potential, stomatal conductance, leaf transpiration and leaf specific hydraulic conductance, there were no differences in midday leaf water potentials. The strong regulation of changes in leaf water potential in Q. suber both diurnally and seasonally is achieved through stomatal closure, which is sensitive to changes in both liquid and vapor phase conductance. This sensitivity allows for optimization of carbon and water resource use without compromising the root-shoot hydraulic link.     

Leaf hydraulic conductance in relation to anatomical and functional traits during Populus tremula leaf ontogeny

Leaf hydraulic conductance (K(leaf)) and several characteristics of hydraulic architecture and physiology were measured during the first 10 weeks of leaf ontogeny in Populus tremula L. saplings growing under control, mild water deficit or elevated temperature conditions. During the initial 3 weeks of leaf ontogeny, most measured characteristics rapidly increased. Thereafter, a gradual decrease in K(leaf) was correlated with a decrease in leaf osmotic potential under all conditions, and with increases in leaf dry mass per area and bulk modulus of elasticity under mild water deficit and control conditions. From about Week 3 onward, K(leaf) was 33% lower in trees subjected to mild water deficit and 33% higher in trees held at an elevated temperature relative to control trees. Mild water deficit and elevated temperature treatment had significant and opposite effects on most of the other characteristics measured. The ontogenetic maximum in K(leaf) was correlated positively with the width of xylem conduits in the midrib, but negatively with the overall width of the midrib xylem, number of lateral ribs, leaf dry mass per area and bulk modulus of elasticity. The ontogenetic maximum in K(leaf) was also correlated positively with the proportion of intercellular spaces and leaf osmotic potential, but negatively with leaf thickness, volume of mesophyll cells and epidermis and number of cells per total mesophyll cell volume, the closest relationships being between leaf osmotic potential and number of cells per total mesophyll cell volume. It was concluded that differences in protoplast traits are more important than differences in xylem or parenchymal cell wall traits in determining the variability in K(leaf) among leaves growing under different environmental conditions.     

Quercus species differ in water and nutrient characteristics in a resource-limited fall-line sandhill habitat

We compared co-occurring mature Quercus laevis Walt. (turkey oak), Q. margaretta Ashe (sand post oak) and Q. incana Bartr. (bluejack oak) trees growing in resource-limited sandhill habitats of the southeastern United States for water and nutrient characteristics. The Quercus spp. differed in their distribution along soil water and nutrient gradients, and in their access to and use of water, even though the study year was wetter than average with no mid-season drought. Quercus laevis had the greatest access to soil water (least negative pre-dawn water potential, psi(pd)) and the most conservative water-use strategy based on its relatively low stomatal conductance (g(s)), high instantaneous water-use efficiency (WUE), least negative midday water potential (psy(md)) and high leaf specific hydraulic conductance (K(L)). Quercus margaretta had the least conservative water-use characteristics, exhibiting relatively high g(s), low instantaneous WUE, most negative psi(md), and low K(L). Quercus margaretta also had a low photosynthetic nitrogen-use efficiency (PNUE), but a high leaf phosphorus concentration. Quercus incana had the poorest access to soil water, but intermediate water-use characteristics and leaf nutrient characteristics more similar to those of Q. laevis. There were no species differences for photosynthesis (A), leaf nitrogen on an area basis, or seasonally integrated WUE (delta13C). Both A and g(s) were positively correlated for each species, but A and g(s) were generally not correlated with psi(pd), psi(md) or delta psi(pd-md). Although we found differences in resource use and resource status among these sandhill Quercus spp., the results are consistent with the interpretation that they are generally drought avoiders. Quercus laevis may have an advantage on xeric ridges because of its greater ability to access soil water and use it more conservatively compared with the other Quercus spp.     

Associations between growth, wood anatomy, carbon isotope discrimination and mortality in a Quercus robur forest

Observations of forest mortality are increasing globally, but relatively little is known regarding the underlying mechanisms driving these events. Tree rings carry physiological signatures that may be used as a tool for retrospective analyses. We capitalized on a local soil water drainage event in 1982 that resulted in increased mortality within a stand of oak trees (Quercus robur), to examine the underlying physiological patterns associated with survival and death in response to soil water limitations. Pre-dawn water potentials showed more negative values for trees in the process of dying compared with those that survived. We used tree rings formed over the 123 years prior to mortality to estimate productivity from basal area increment (BAI, mm(2)), multiple xylem hydraulic parameters via anatomical measurements and crown-level gas exchange via carbon isotope discrimination (Δ, ‰). Oaks that died had significantly higher BAI values than trees that survived until the drainage event, after which the BAI of trees that died declined dramatically. Hydraulic diameter and conductivity of vessels in trees that died were higher than in surviving trees until the last 5 years prior to mortality, at which time both groups had similar values. Trees that died had consistently lower Δ values than trees that survived. Therefore, tree mortality in this stand was associated with physiological differences prior to the onset of soil water reduction. We propose that trees that died may have been hydraulically underbuilt for dry conditions, which predisposes them to severe hydraulic constraints and subsequent mortality. Measurements of above-ground/below-ground dry mass partitioning will be critical to future tests of this hypothesis. Based on these results, it is probable that pedunculate oak trees will experience greater future mortality if climate changes cause more severe droughts than the trees have experienced previously.     

Variation in drought resistance, drought acclimation and water conservation in four willow cultivars used for biomass production

Growth and water-use parameters of four willow (Salix spp.) clones grown in a moderate drought regime or with ample water supply were determined to characterize their water-use efficiency, drought resistance and capacity for drought acclimation. At the end of the 10-week, outdoor pot experiment, clonal differences were observed in: (1) water-use efficiency of aboveground biomass production (WUE); (2) resistance to xylem cavitation; and (3) stomatal conductance to leaf-specific, whole-plant hydraulic conductance ratio (g(st)/K(P); an indicator of water balance). Across clones and regimes, WUE was positively correlated with the assimilation rate to stomatal conductance ratio (A/g(st)), a measure of instantaneous water-use efficiency. Both of these water-use efficiency indicators were generally higher in drought-treated trees compared with well-watered trees. However, the between-treatment differences in (shoot-based) WUE were smaller than expected, considering the differences in A/g(st) for two of the clones, possibly because plants reallocated dry mass from shoots to roots when subject to drought. Higher root hydraulic conductance to shoot hydraulic conductance ratios (K(R)/K(S)) during drought supports this hypothesis. The same clones were also the most sensitive to xylem cavitation and, accordingly, showed the strongest reduction in g(st)/K(P) in response to drought. Drought acclimation was manifested in decreased g(st), g(st)/K(P), osmotic potential and leaf area to vessel internal cross-sectional area ratio, and increased K(R), K(P) and WUE. Increased resistance to stem xylem cavitation in response to drought was observed in only one clone. It is concluded that WUE and drought resistance traits are inter-linked and that both may be enhanced by selection and breeding.     

Boxelder water sources and physiology at perennial and ephemeral stream sites in Arizona

To assess the influence of stream water on leaf gas exchange and water potential in different sized boxelder trees (Acer negundo L.), we compared these characteristics in trees growing beside a perennial stream and a nearby ephemeral stream in a montane-riparian forest in northern Arizona. Patterns of tree water use were quantified by stable isotope analysis (delta(18)O). Physiological characteristics were similar for large and small trees. Similarity between sites in predawn and daytime water potentials and xylem delta(18)O indicated that stream water was not a physiologically important water source. Seasonal and site variations in light-saturated net photosynthetic rate were significantly related to leaf-to-air vapor pressure deficit (r = -0.691) and foliar nitrogen concentration (r = 0.388). Although deep water was the dominant water source, surface soil water was utilized following precipitation, especially by small trees. We conclude that net carbon gain and severity of water stress are only weakly coupled to stream water availability.     

Water infiltration and hydraulic conductivity in sandy cambisols: impacts of forest transformation on soil hydrological properties

Soil hydrological properties like infiltration capacity and hydraulic conductivity have important consequences for hydrological properties of soils in river catchments and for flood risk prevention. They are dynamic properties due to varying land use management practices. The objective of this study was to characterize the variation of infiltration capacity, hydraulic conductivity and soil organoprofile development on forest sites with comparable geological substrate, soil type and climatic conditions, but different stand ages and tree species in terms of the effects of forest transformation upon soil hydrological properties. The Kahlenberg forest area (50 km northeast of Berlin in the German northeastern lowlands) under investigation contains stands of Scots pine (Pinus sylvestris) and European beech (Fagus sylvatica) of different age structures forming a transformation chronosequence from pure Scots pine stands towards pure European beech stands. The water infiltration capacity and hydraulic conductivity (K) of the investigated sandy-textured soils are low and very few macropores exist. Additionally these pores are marked by poor connectivity and therefore do not have any significant effect on water infiltration rate. Moreover, water infiltration in these soils is impeded by their hydrophobic properties. Along the experimental chronosequence of forest transformation, the thickness of the forest floor layer decreases due to enhanced decomposition and humification intensities. By contrast, the thickness of the humous topsoil increases. Presumably, changes in soil organic matter storage and quality caused by the management practice of forest transformation affect the persistence and degree of water repellency in the soil, which in turn influences the hydraulic properties of the experimental soils. The results indicate clearly that soils play a crucial role for water retention and therefore, in overland flow prevention. There is a need to have more awareness on the intimate link between the land use and soil properties and their possible effects on flooding.     

Water loss regulation in mature Hevea brasiliensis: effects of intermittent drought in the rainy season and hydraulic regulation

Effects of soil and atmospheric drought on whole-tree transpiration (E(T)), leaf water potential (Ψ(L)) and whole-tree hydraulic conductance (K(T)) were investigated in mature rubber trees (Hevea brasiliensis, clone RRIM 600) during the full canopy stage in the rainy season in a drought-prone area of northeast Thailand. Under well-watered soil conditions, transpiration was tightly regulated in response to high evaporative demand, i.e., above reference evapotranspiration (ET(0)) ~2.2 mm day(-1) or maximum vapor pressure deficit ~1.8 kPa. When the trees experienced intermittent soil drought E(T) decreased sharply when relative extractable water in the top soil was?相似文献   

Can the nitrogenous composition of xylem sap be used to assess salinity stress in Casuarina glauca?

It is predicted that dryland salinity will affect up to 17 Mha of the Australian landscape by 2050, and therefore, monitoring the health of tree plantings and remnant native vegetation in saline areas is increasingly important. Casuarina glauca Sieber ex Spreng. has considerable salinity tolerance and is commonly planted in areas with a shallow, saline water table. To evaluate the potential of using the nitrogenous composition of xylem sap to assess salinity stress in C. glauca, the responses of trees grown with various soil salinities in a greenhouse were compared with those of trees growing in field plots with different water table depths and groundwater salinities. In the greenhouse, increasing soil salinity led to increased allocation of nitrogen (N) to proline and arginine in both stem and root xylem sap, with coincident decreases in citrulline and asparagine. Although the field plots were ranked as increasingly saline-based on ground water salinity and depth-only the allocation of N to citrulline differed significantly between the field plots. Within each plot, temporal variation in the composition of the xylem sap was related to rainfall, rainfall infiltration and soil salinity. Periods of low rainfall and infiltration and higher soil salinity corresponded with increased allocation of N to proline and arginine in the xylem sap. The allocation of N to citrulline and asparagine increased following rainfall events where rain was calculated to have infiltrated sufficiently to decrease soil salinity. The relationship between nitrogenous composition of the xylem sap of C. glauca and soil salinity indicates that the analysis of xylem sap is an effective method for assessing changes in salinity stress in trees at a particular site over time. However, the composition of the xylem sap proved less useful as a comparative index of salinity stress in trees growing at different sites.     

Transpiration and water relations of poplar trees growing close to the water table

Sap flow was measured on five branches of two poplar (Populus trichocarpa Torr. & A. Gray x P. tacamahaca L.) trees from June to September 1994 in the south of England with stem-surface, heat balance gauges, and was scaled up to estimate transpiration from single trees on the basis of leaf area. On six days, stomatal conductance and plant water potential were measured simultaneously with a porometer and pressure chamber, respectively. The effects of solar radiation (S), vapor pressure deficit (D) and stomatal conductance on transpiration were evaluated. Sap flow per unit leaf area (F(a)) was closely related to the time course of demand attributable to S and D throughout the season, and only slightly affected by the water content of the top 120 cm of soil. Although F(a) increased linearly at low values of D, it showed a plateau with increases in D above 1.2 kPa. The canopy coupling coefficient (1 - Omega) ranged from 0.48 to 0.78 with a mean of 0.65 +/- 0.01, indicating that transpiration was controlled more by stomatal conductance than by incident radiation. The seasonal pattern of tree water loss followed potential evaporation with a peak in late June or early July. On bright days, daily transpiration over the projected crown area was 3.6 mm early in the season, 3.8 mm in mid-season, and 2.7 mm late in the season. The water balance of the system indicated that poplar trees took 15-60% of water transpired from groundwater, with the proportion increasing as the soil in the unsaturated zone dried out. Access to the water table resulted in high predawn water potentials throughout the season. Estimated hydraulic resistance to water flow in the soil-tree system was in the range of 1.5 to 1.93 x 10(6) MPa s m(-3).     

Water-use strategies in two co-occurring Mediterranean evergreen oaks: surviving the summer drought

In the Mediterranean evergreen oak woodlands of southern Portugal, the main tree species are Quercus ilex ssp. rotundifolia Lam. (holm oak) and Quercus suber L. (cork oak). We studied a savannah-type woodland where these species coexist, with the aim of better understanding the mechanisms of tree adaptation to seasonal drought. In both species, seasonal variations in transpiration and predawn leaf water potential showed a maximum in spring followed by a decline through the rainless summer and a recovery with autumn rainfall. Although the observed decrease in predawn leaf water potential in summer indicates soil water depletion, trees maintained transpiration rates above 0.7 mm day(-1) during the summer drought. By that time, more than 70% of the transpired water was being taken from groundwater sources. The daily fluctuations in soil water content suggest that some root uptake of groundwater was mediated through the upper soil layers by hydraulic lift. During the dry season, Q. ilex maintained higher predawn leaf water potentials, canopy conductances and transpiration rates than Q. suber. The higher water status of Q. ilex was likely associated with their deeper root systems compared with Q. suber. Whole-tree hydraulic conductance and minimum midday leaf water potential were lower in Q. ilex, indicating that Q. ilex was more tolerant to drought than Q. suber. Overall, Q. ilex seemed to have more effective drought avoidance and drought tolerance mechanisms than Q. suber.     

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

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