Whole-tree Hydraulic Conductance Research Articles

The increase of leaf water potential and whole-tree hydraulic conductance promotes canopy conductance and transpiration of Pinus tabulaeformis during soil droughts

The increase of leaf water potential and whole-tree hydraulic conductance promotes canopy conductance and transpiration of Pinus tabulaeformis during soil droughts

Coordination between liquid and gas phase conductance enables survival during water stress in an isohydrodynamic Eucalyptus cultivar

The regulation on transpiration and water status is critical for tree survival during drought. Eucalyptus is introduced and widely planted in southern China for timber production. Eucalyptus plantations are expanding into rugged terrains with shallow soils and soil water capacitance. With the ongoing decline in precipitation and the typical monsoon climate in this region, the trees suffer regular seasonal drought, yet their response mechanisms to drought remain obscure. We examined leaf water status, stomatal and hydraulic behavior, and whole-tree transpiration of a wildly planted eucalyptus cultivar, Eucalyptus urophylla × Eucalyptus grandis, across seasons with different water availability in order to identify its response strategy to drought. During the study period, both predawn (ΨPD) and midday (ΨMD) leaf water potential varied considerably, but the water potential gradient between the roots and leaves (ΔΨS-L), measured as the difference between ΨPD and ΨMD, remained constant, indicating an isohydrodynamic regulation on water status under drought. Despite strong stomatal down regulation on transpiration in response to increasing evaporative demand, leaf water potential declined to levels approaching -2 MPa at midday in the driest months, inducing more than 60% loss of xylem hydraulic conductance. Both stomatal and whole-tree hydraulic conductance linearly declined with increasing drought intensity. Whole-tree hydraulic conductance was exponentially correlated with the sensitivity of stomatal conductance to VPD. These results demonstrated that stomatal and hydraulic regulation on transpiration of eucalyptus was coupled through tight correlation between stomatal response to evaporative demand and whole-tree hydraulic conductance. This coordination between liquid and gas phase conductance found in this study was helpful to keep a balance between allowable maximum transpiration and hydraulic safety of eucalyptus under varying soil and atmosphere moisture conditions.

Functional Relationships Between Branch and Stem Wood Density for Temperate Tree Species in North America

Wood density is strongly related to key aspects of tree physiological performance. While many studies have examined wood density in different parts of trees for a variety of reasons, there are very few studies that have compared within-tree density variation across many trees, of many species, drawn from a large geographic area. Here, a large data set representing trees of 78 species/genera in the Eastern United States was compiled and analyzed to explore branch to main stem wood basic density relationships. The expectation was that differences in stem versus branch wood density among trees would be due to both genetic constraints and plastic responses in wood properties, due to tree growth responses to external environments. The results show a wide tree-to-tree variation in average branch density, relative to main stem density. However, there was a general pattern for overstory tree species to have high tree branch density relative to stem density, at lower stem densities, and a declining branch to stem wood density ratio as stem density increased. Evergreen gymnosperms showed the strongest change in branch to stem wood density ratios over the range of stem wood densities and deciduous angiosperms the least; deciduous gymnosperms showed an intermediate pattern, but with generally higher branch- than stem- wood densities. More cold-hearty, shade-tolerant / drought-intolerant, evergreen gymnosperms, growing at higher latitudes, showed higher branch to stem density ratios than more shade-intolerant / drought tolerant evergreen gymnosperms growing at lower latitudes. Across all trees, canopy position had a significant influence on branch to stem density relationships, with higher branch to stem density ratios for canopy dominant trees and successively lower branch to stem density ratios for trees in successively inferior canopy positions (in terms of light availability). Understory tree species, which remain in the forest understory at maximum height, showed generally lower branch than stem densities over a wide range of stem densities. The results suggested that tradeoffs between mechanical safety and whole-tree hydraulic conductance are driving within-tree differences in wood density and highlighted the need for more detailed examinations of within-tree density variation at the whole-tree level.

Limited stomatal regulation of the largest-size class of Dryobalanops aromatica in a Bornean tropical rainforest in response to artificial soil moisture reduction.

The physiological response of trees to drought is crucial for understanding the risk of mortality and its feedbacks to climate under the increase in droughts due to climate change, especially for the largest trees in tropical rainforests because of their large contribution to total carbon storage and water use. We determined the response of the mean canopy stomatal conductance per unit leaf area (gs) and whole-tree hydraulic conductance (Gp) of the largest individuals (38-53m in height) of a typical canopy tree species in a Bornean tropical rainforest, Dryobalanops aromatica C.F.Gaertn., to soil moisture reduction by a 4-month rainfall exclusion experiment (REE) based on the measurements of sap flux and leaf water potentials at midday and dawn. In the mesic condition, the gs at vapor pressure deficit (D) = 1kPa (gsref) was small compared with the reported values in various biomes. The sensitivity of gs to D (m) at a given gsref (m/gsref) was ≥ 0.6 irrespective of soil moisture conditions, indicating intrinsically sensitive stomatal control with increasing D. The REE caused greater soil drought and decreased the mean leaf water potentials at midday and dawn to the more negative values than the control under the relatively dry conditions due to natural reduction in rainfall. However, the REE did not cause a greater decrease in gs nor any clear alteration in the sensitivity of gs to D compared with the control, and induced greater decreases in Gp during REE than the control. Thus, though the small gs and the sensitive stomatal response to D indicate the water saving characteristics of the studied trees under usual mesic conditions, their limited stomatal regulation in response to soil drought by REE and the resulting decline in Gp might suggest a poor resistance to the unusually severe drought expected in the future.

Tree Water Use, Water Use Efficiency, and Carbon Isotope Discrimination in Relation to Growth Potential in Populus deltoides and Hybrids under Field Conditions

We explored the relationship between tree growth, water use, and related hydraulic traits in Populus deltoides Bartr. ex Marsh.and hybrid clones, to examine potential trade-offs between growth and water use efficiency. Nine genotypes, six P. deltoides and three hybrid clones, that represented genotypes with high (Group H), intermediate (Group I), and low (Group L) growth performance were selected for study, based on year-two standing stem biomass in a replicated field trial. In year four, tree growth, transpiration (Et), canopy stomatal conductance (Gs), whole-tree hydraulic conductance (Gp), and carbon isotope discrimination (Δ13C) were measured. Tree sap flux was measured continuously using thermal dissipation probes. We hypothesized that Group H genotypes would have increased growth efficiency (GE), increased water use efficiency of production (WUEp, woody biomass growth/Et), lower Δ13C, and greater Gp than slower growing genotypes. Tree GE increased with relative growth rate (RGR), and mean GE in Group H was significantly greater than L, but not I. Tree WUEp ranged between 1.7 and 3.9 kg biomass m3 H2O−1, which increased with RGR. At similar levels of Et, WUEp was significantly greater in Group H (2.45 ± 0.20 kg m−3), compared to I (2.03 ± 0.18 kg m−3) or L (1.72 ± 0.23 kg m−3). Leaf and wood Δ13C scaled positively with stem biomass growth but was not correlated with WUEp. However, at a similar biomass increment, clones in Group H and I had significantly lower leaf Δ13C than Group L. Similarly, Group H clones had a significantly lower wood Δ13C than Group L, supporting our hypothesis of increased WUE in larger trees. Tree physiological and hydraulic traits partially explain differences in WUEp and Δ13C, and suggest that clone selection and management activities that increase tree biomass production will likely increase tree and stand WUE. However, more research is needed to discern the underlying hydraulic mechanisms responsible for the higher WUE exhibited by large trees and distinct clones.

Excessive nitrogen impairs hydraulics, limits photosynthesis, and alters the metabolic composition of almond trees

Horticulture nitrogen (N) runoffs are major environmental and health concerns, but current farming practices cannot detect ineffective N applications. Hence, we set to recognize high N conditions and characterize their effects on the physiology of almond trees grown in drainage lysimeters. Water and nutrients mass balances exhibited that N benefitted almond trees in a limited range (below 60 mg N L−1 in irrigation), while higher N conditions (over a 100 mg N L−1) reduced evapotranspiration (ET) by 50% and inherently constrained N uptake. Respectively, whole-tree hydraulic conductance reduced by 37%, and photosynthesis by 17%, which implied that high N concentrations could damage trees. Through gas-chromatography, we realized that high N conditions also affected components of the citric acid cycle (TCA) and carbohydrates availability. Such changes in the metabolic composition of roots and leaves probably interfered with N assimilation and respiration. It also determined the proportions between N and starch in almond leaves, which formed a new index (N:ST) that starts at 0.4 in N deficiency and reaches 0.6–0.8 in optimal N conditions. Importantly, this index continues to increase in higher N conditions (as starch reduces) and essentially indicates to excessive N applications when it exceeds 1.1.

Using sap flow to measure whole-tree hydraulic conductance loss in response to drought

Xylem vulnerability to cavitation is an important physiological trait that strongly influences how plants respond to drought. There are several techniques available to measure branch-level xylem vulnerability to cavitation, however these are often time and resource consuming and might not always reflect the hydraulic bottlenecks on the soil-canopy continuum. These characteristics restrict the availability of tissue specific hydraulic traits and its use in large-scale modelling. In this study we propose a sap flow-based approach to estimate hydraulic parameters that describe the whole-tree conductance loss in response to drought. The hydraulic parameters estimated by our whole-tree approach are close to the branch-level parameters measured with the traditional bench drying technique. Consequently, the hydraulic parameters derived from both approaches produce similar model predictions of plant responses to soil and atmospheric drought. This indicates a coordinated loss of conductance among the different tissues that compose the soil-canopy hydraulic continuum of the studied species (Drimys brasiliensis). Our approach provides an alternative and easier way to characterize whole-plant hydraulic response to drought and could be useful to produce plant and ecosystem-level hydraulic parameters to be used in large-scale vegetation models.

Dry and hot: the hydraulic consequences of a climate change-type drought for Amazonian trees.

How plants respond physiologically to leaf warming and low water availability may determine how they will perform under future climate change. In 2015-2016, an unprecedented drought occurred across Amazonia with record-breaking high temperatures and low soil moisture, offering a unique opportunity to evaluate the performances of Amazonian trees to a severe climatic event. We quantified the responses of leaf water potential, sap velocity, whole-tree hydraulic conductance (Kwt), turgor loss and xylem embolism, during and after the 2015-2016 El Niño for five canopy-tree species. Leaf/xylem safety margins (SMs), sap velocity and Kwt showed a sharp drop during warm periods. SMs were negatively correlated with vapour pressure deficit, but had no significant relationship with soil water storage. Based on our calculations of canopy stomatal and xylem resistances, the decrease in sap velocity and Kwt was due to a combination of xylem cavitation and stomatal closure. Our results suggest that warm droughts greatly amplify the degree of trees' physiological stress and can lead to mortality. Given the extreme nature of the 2015-2016 El Niño and that temperatures are predicted to increase, this work can serve as a case study of the possible impact climate warming can have on tropical trees.This article is part of a discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

Culm Age and Rhizome Affects Night-Time Water Recharge in the Bamboo Phyllostachys pubescens.

Bamboo species—the only herbaceous trees—have unique structural and physiological characteristics that differ from those of other tree taxa. However, the role of night-time water use in bamboo is poorly understood and has rarely been investigated. We studied the day- and night-time sap flow response to culm age and rhizome structure in three age levels (juvenile, mature, and senescent) of Phyllostachys pubescens growing in the Nankun Mountain Natural Reserve, South China. We found that sap flow density and whole-tree hydraulic conductance decreased with culm age. After cutting of rhizome, the day-time sap flow and night-time water recharge decreased obviously. In addition, night-time water recharge accounted for the largest proportion (up to 30%) of total daily transpiration in normal senescent bamboos. Therefore, our study indicates that the connected rhizome system and night-time water recharge played a significant role in water compensation during the day and at night in bamboos. Night-time water recharge is especially critical to senescent bamboos, given their weaker transpiration due to the lower whole-tree hydraulic conductance, and consequently, they are more dependent on night-time water recharge for fulfilling their whole-day water consumption needs.

Loblolly Pine Productivity and Water Relations in Response to Throughfall Reduction and Fertilizer Application on a Poorly Drained Site in Northern Florida

Loblolly pine (Pinus taeda L.) forests are of great ecological and economic value in the southeastern United States, where nutrient availability frequently limits productivity. The impact of fertilizer application on the growth and water relations of loblolly pine has been investigated by numerous studies; however, few field experiments have examined the effects of drought. Drought is of particular interest due to the potential for climate change to alter soil water availability. In this study, we investigated the impact of fertilizer application and a 30% reduction in throughfall on loblolly pine productivity, transpiration, hydraulic conductance, and stomatal conductance. The study was installed in a ten-year-old loblolly pine plantation on a somewhat poorly drained site in northern Florida. Throughfall reduction did not impact tree productivity or water relations of the trees. This lack of response was attributed to abundant rainfall and the ability of trees to access the shallow water table at this site. Fertilizer application increased basal area production by 20% and maximum leaf area index by 0.5 m2∙m−2, but it did not affect whole-tree hydraulic conductance or the sensitivity of stomatal conductance to vapor pressure deficit. During the spring, when leaf area and vapor pressure deficit were high, the fertilizer-only treatment increased monthly transpiration by 17% when compared to the control. This relationship, however, was not significant during the rest of the year.

Loss of whole-tree hydraulic conductance during severe drought and multi-year forest die-off

Understanding the pathways through which drought stress kills woody vegetation can improve projections of the impacts of climate change on ecosystems and carbon-cycle feedbacks. Continuous in situ measurements of whole trees during drought and as trees die hold promise to illuminate physiological pathways but are relatively rare. We monitored leaf characteristics, water use efficiency, water potentials, branch hydraulic conductivity, soil moisture, meteorological variables, and sap flux on mature healthy and sudden aspen decline-affected (SAD) trembling aspen (Populus tremuloides) ramets over two growing seasons, including a severe summer drought. We calculated daily estimates of whole-ramet hydraulic conductance and modeled whole-ramet assimilation. Healthy ramets experienced rapid declines of whole-ramet conductance during the severe drought, providing an analog for what likely occurred during the previous drought that induced SAD. Even in wetter periods, SAD-affected ramets exhibited fivefold lower whole-ramet hydraulic conductance and sevenfold lower assimilation than counterpart healthy ramets, mediated by changes in leaf area, water use efficiency, and embolism. Extant differences between healthy and SAD ramets reveal that ongoing multi-year forest die-off is primarily driven by loss of whole-ramet hydraulic capability, which in turn limits assimilation capacity. Branch-level measurements largely captured whole-plant hydraulic limitations during drought and mortality, but whole-plant measurements revealed a potential role of other losses in the hydraulic continuum. Our results highlight the importance of a whole-tree perspective in assessing physiological pathways to tree mortality and indicate that the effects of mortality on these forests' assimilation and productivity are larger than expected based on canopy leaf area differences.

Drought‐induced defoliation and long periods of near‐zero gas exchange play a key role in accentuating metabolic decline of Scots pine

Drought-induced defoliation has recently been associated with the depletion of carbon reserves and increased mortality risk in Scots pine (Pinus sylvestris). We hypothesize that defoliated individuals are more sensitive to drought, implying that potentially higher gas exchange (per unit of leaf area) during wet periods may not compensate for their reduced photosynthetic area. We measured sap flow, needle water potentials and whole-tree hydraulic conductance to analyse the drought responses of co-occurring defoliated and nondefoliated Scots pines in northeast Spain during typical (2010) and extreme (2011) drought conditions. Defoliated Scots pines showed higher sap flow per unit leaf area during spring, but were more sensitive to summer drought, relative to nondefoliated pines. This pattern was associated with a steeper decline in soil-to-leaf hydraulic conductance with drought and an enhanced sensitivity of canopy conductance to soil water availability. Near-homeostasis in midday water potentials was observed across years and defoliation classes, with minimum values of -2.5MPa. Enhanced sensitivity to drought and prolonged periods of near-zero gas exchange were consistent with low levels of carbohydrate reserves in defoliated trees. Our results support the critical links between defoliation, water and carbon availability, and their key roles in determining tree survival and recovery under drought.

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 < 0.4. The midday leaf water potential (Ψ(md)) on sunny days did not change as a function of soil drought and remained stable at approximately - 1.95 MPa, i.e., displaying isohydric behavior. The decrease in E(T) was mainly due to the change in K(T). K(T) remained constant over a wide range of environmental conditions and decreased sharply at low soil water availability. A simple hydraulic model incorporating critical minimum water potential and the response of whole-tree hydraulic conductance to relative extractable water correctly simulated patterns of transpiration over 6 months. We conclude that an explicit and simplified framework of hydraulic limitation hypothesis was sufficient to describe water use regulation of a mature rubber tree stand in water-limited conditions. Given the complexity of constraints in the soil-plant-atmosphere pathway, our results confirm the relevance of this approach to synthesize the overall behavior of trees under drought.

Comparative hydraulic architecture of tropical tree species representing a range of successional stages and wood density

Plant hydraulic architecture (PHA) has been linked to water transport sufficiency, photosynthetic rates, growth form and attendant carbon allocation. Despite its influence on traits central to conferring an overall competitive advantage in a given environment, few studies have examined whether key aspects of PHA are indicative of successional stage, especially within mature individuals. While it is well established that wood density (WD) tends to be lower in early versus late successional tree species, and that WD can influence other aspects of PHA, the interaction of WD, successional stage and the consequent implications for PHA have not been sufficiently explored. Here, we studied differences in PHA at the scales of wood anatomy to whole-tree hydraulic conductance in species in early versus late successional Panamanian tropical forests. Although the trunk WD was indistinguishable between the successional groups, the branch WD was lower in the early successional species. Across all species, WD correlated negatively with vessel diameter and positively with vessel packing density. The ratio of branch:trunk vessel diameter, branch sap flux and whole-tree leaf-specific conductance scaled negatively with branch WD across species. Pioneer species showed greater sap flux in branches than in trunks and a greater leaf-specific hydraulic conductance, suggesting that pioneer species can move greater quantities of water at a given tension gradient. In combination with the greater water storage capacitance associated with lower WD, these results suggest these pioneer species can save on the carbon expenditure needed to build safer xylem and instead allow more carbon to be allocated to rapid growth.

Whole-tree water relations of co-occurring mature Pinus palustris and Pinus elliottii var.elliottii

The natural range of longleaf pine ( Pinus palustris P. Mill.) and slash pine ( Pinus elliottii var. elliottii Engelm.) includes most of the southeastern US Coastal Plain, and there is now considerable interest in using these species for ecological forestry, restoration, and carbon sequestration. It is therefore surprising that little information is currently available concerning differences in their ecological water relations in natural stands. In this study, we compared water use, stomatal conductance at the crown scale (Gcrown), and whole-tree hydraulic conductance of mature pine trees growing in a naturally regenerated mixed stand on a flatwoods site in north-central Florida. We found remarkable similarities between longleaf and slash pine in stored water use, nocturnal transpiration, and whole-tree hydraulic conductance. Mean daily transpiration rate was higher for slash than for longleaf pine, averaging 39 and 26 L·tree–1, respectively. This difference was determined by variations in tree leaf area. Slash pine had 60% more leaf area per unit basal sapwood area than longleaf pine, but the larger plasticity of longleaf pine stomatal regulation partially compensated for leaf area differences: longleaf pine had higher Gcrownon days with high volumetric water content (θv) but this was reduced to similar or even lower values than for slash pine on days with low θv. There was no species difference in the sensitivity of Gcrownto increasing vapor pressure deficit.

Variable conductivity and embolism in roots and branches of four contrasting tree species and their impacts on whole-plant hydraulic performance under future atmospheric CO2 concentration

Anatomical and physiological acclimation to water stress of the tree hydraulic system involves trade-offs between maintenance of stomatal conductance and loss of hydraulic conductivity, with short-term impacts on photosynthesis and long-term consequences to survival and growth. Here, we study the role of variations in root and branch maximum hydraulic specific conductivity (k(s-max)) under high and low soil moisture in determining whole-tree hydraulic conductance (K(tree)) and in mediating stomatal control of gas exchange in four contrasting tree species growing under ambient and elevated CO₂ (CO₂(a) and CO₂(e)). We hypothesized that K(tree) would adjust to CO₂(e) through an increase in root and branch k(s-max) in response to anatomical adjustments. However, physiological changes observed under CO₂(e) were not clearly related to structural change in the xylem of any of the species. The only large effect of CO₂(e) occurred in branches of Liquidambar styraciflua L. and Cornus florida L. where an increase in k(s-max) and a decrease in xylem resistance to embolism (-P₅₀) were measured. Across species, embolism in roots explained the loss of K(tree) and therefore indirectly constituted a hydraulic signal involved in stomatal regulation and in the reduction of G(s-ref), the sap-flux-scaled mean canopy stomatal conductance at a reference vapour pressure deficit of 1 kPa. Across roots and branches, the increase in k(s-max) was associated with a decrease in -P₅₀, a consequence of structural acclimation such as larger conduits, lower pit resistance and lower wood density. Across species, treatment-induced changes in K(tree) translated to similar variation in G(s-ref). However, the relationship between G(s-ref) and K(tree) under CO₂(a) was steeper than under CO₂(e), indicating that CO₂(e) trees have lower G(s-ref) at a given K(tree) than CO₂(a) trees. Under high soil moisture, CO₂(e) greatly reduced G(s-ref). Under low soil moisture, CO₂(e) reduced G(s-ref) of only L. styraciflua and Ulmus alata. In some species, higher xylem dysfunction under CO₂(e) might impact tree performance in a future climate when increased evaporative demand could cause a greater loss of hydraulic function. The results contributed to our knowledge of the physiological and anatomical mechanisms underpinning the responses of tree species to drought and more generally to global change.

Stable annual pattern of water use by Acacia tortilis in Sahelian Africa

Water use by mature trees of Acacia tortilis (Forsk.) Hayne ssp. raddiana (Savi) Brenan var. raddiana growing in the northern Sahel was continuously recorded over 4 years. Water use was estimated from xylem sap flow measured by transient heat dissipation. Concurrently, cambial growth, canopy phenology, leaf water potential, climatic conditions and soil water availability (SWA) were monitored. In addition to the variation attributable to interannual variation in rainfall, SWA was increased by irrigation during one wet season. The wet season lasted from July to September, and annual rainfall ranged between 146 and 367 mm. The annual amount and pattern of tree water use were stable from year-to-year despite interannual and seasonal variations in SWA in the upper soil layers. Acacia tortilis transpired readily throughout the year, except for one month during the dry season when defoliation was at a maximum. Maximum water use of about 23 l (dm sapwood area)(-2) day(-1) was recorded at the end of the wet season. While trees retained foliage in the dry season, the decline in water use was modest at around 30%. Variation in predawn leaf water potential indicated that the trees were subject to soil water constraint. The rapid depletion of water in the uppermost soil layers after the wet season implies that there was extensive use of water from deep soil layers. The deep soil profile revealed (1) the existence of living roots at 25 m and (2) that the availability of soil water was low (-1.6 MPa) down to the water table at a depth of 31 m. However, transpiration was recorded at a predawn leaf water potential of -2.0 MPa, indicating that the trees used water from both intermediary soil layers and the water table. During the full canopy stage, mean values of whole-tree hydraulic conductance were similar in the wet and dry seasons. We propose that the stability of water use at the seasonal and annual scales resulted from a combination of features, including an extensive rooting habit related to deep water availability and an effective regulation of canopy conductance. Despite a limited effect on tree water use, irrigation during the wet season sharply increased predawn leaf water potential and cambial growth of trunks and branches.

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.

Tree height and age-related decline in growth in Scots pine (Pinus sylvestris L.)

Growth and seasonal water use was measured amongst trees growing in an old growth Scots pine forest in the Scottish Highlands. Three sites which differed in their recent management history and contained old and naturally regenerated young trees growing together were monitored in the field. Our results showed a clear decrease in growth efficiency with age, from values of around 0.25 kg m(-2) leaves year(-1) in approximately 25-year-old trees to less than 0.1 kg m(-2) leaves year(-1) in trees over 200 years old. When the old trees in one of the field sites were released from competition by thinning, their growth efficiency reverted to that of coexisting young trees, indicating that the decline in growth was reversible. This is consistent with the results of a parallel study showing that cambial age had no effect on the physiology or growth of grafted seedlings originating from the same population studied here (Mencuccini et al. 2005). Our detailed study of tree water use in the field showed an overall decrease in whole-tree hydraulic conductance and stomatal canopy conductance with tree height in the unthinned stands, in agreement with the hydraulic limitation hypothesis. However, the effect of this reduction in hydraulic efficiency on growth was comparatively small, and old trees also showed consistently lower nitrogen concentrations in needles, suggesting that hydraulic and nutritional factors combined to produce the decline in growth efficiency with age observed in the studied populations.

Comparative water use by the riparian trees Melaleuca argentea and Corymbia bella in the wet-dry tropics of northern Australia

We examined sources of water and daily and seasonal water use patterns in two riparian tree species occupying contrasting niches within riparian zones throughout the wet-dry tropics of northern Australia: Corymbia bella Hill and Johnson is found along the top of the levee banks and Melaleuca argentea W. Fitzg. is restricted to riversides. Patterns of tree water use (sap flow) and leaf water potential were examined in four trees of each species at three locations along the Daly River in the Northern Territory. Predawn leaf water potential was higher than -0.5 MPa throughout the dry season in both species, but was lower at the end of the dry season than at the beginning of the dry season. Contrary to expectations, predawn leaf water potential was lower in M. argentea trees along the river than in C. bella trees along the levees. In contrast, midday leaf water potential was lower in the C. bella trees than in M. argentea trees. There were no seasonal differences in tree water use in either species. Daily water use was lower in M. argentea trees than in C. bella trees. Whole-tree hydraulic conductance, estimated from the slope of the relationship between leaf water potential and sap flow, did not differ between species. Xylem deuterium concentrations indicated that M. argentea trees along the riverbank were principally reliant on river water or shallow groundwater, whereas C. bella trees along the levee were reliant solely on soil water reserves. This study demonstrated strong gradients of tree water use within tropical riparian communities, with implications for estimating riparian water use requirements and for the management of groundwater resources.