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.     

Coast live oak, Quercus agrifolia, susceptibility and response to goldspotted oak borer, Agrilus auroguttatus, injury in southern California

Oak mortality is often associated with a complex of decline factors. We describe the morphological and physiological responses of coast live oak, Quercus agrifolia Née, in California to an invasive insect, the goldspotted oak borer (GSOB), Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), and evaluate drought as a potential inciting factor. Morphological traits of 356 trees were assessed and physiological traits of 70 of these were monitored intensively over one growing season. Morphological characteristics of tree health included crown thinning and dieback; bole staining resulting from larval feeding; density of GSOB adult exit holes; and holes caused by woodpecker feeding. These characteristics were used to rank GSOB infestation/injury into four classes, and taken together, they explained 87% of the variation in a principal component analysis. Drought stress on various size/age and infestation classes of Q. agrifolia was measured by assessing branchlet pre-dawn and solar noon xylem water potential, leaf cell turgor potential, and water use efficiency over one growing season. Both morphological and physiological traits were highly variable in mature and old growth trees. Early summer plant water status (branchlet xylem water potential and water use efficiency) was similar between uninfested and newly colonized trees, suggesting that GSOB are not pre-selecting drought-stressed Q. agrifolia for oviposition. By late summer, leaf water and cell turgor potentials were lower in infested than in uninfested mature trees, suggesting that GSOB infestation causes drought stress in these trees. Among the tree size/age classes, infested old growth trees exhibited the greatest change in water use efficiency over the growing season, and showed greater morphological injury symptoms of decline than infested mature trees. Morphological attributes of decline in Q. agrifolia associated with GSOB were correlated weakly with increasing physiological drought stress among infestation classes of trees. We propose that the collection of morphological responses of Q. agrifolia to GSOB described here can be used to monitor the future expansion of the GSOB distribution as well as the GSOB-induced decline of Q. agrifolia in California.     

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.     

Shrub species affect distinctively the functioning of scattered Quercus ilex trees in Mediterranean open woodlands

A low tree stand density has been showed as necessary to thrive with summer drought in semiarid Mediterranean open woodlands. Shrub encroachment of these open woodlands is currently recommended to guarantee the persistence of the system, due to the nursery effect of shrubs on tree seedling. However, the increase in abundance and cover of a shrub understory in these water limited woodlands could bring consequences to tree overstory functioning. The present study analyzes the physiological status of scattered Quercus ilex L. trees in paired adjacent plots with and without the presence of a shrubby understory in CW Spain. Two contrasting shrub strategies were addressed in order to take into account possible species-specific effects: a dense-shallow rooting shrub (Cistus landanifer L.) and a sparse-deep rooting shrub (Retama sphaerocarpa (L.) Boiss). Leaf water potential (at predawn and midday), leaf gas exchange parameters (net photosynthetic rate and stomatal conductance), leaf nitrogen content and chlorophyll fluorescence transients (maximum photochemical efficiency and performance index, sensuStrasser et al., 2004) were measured during three consecutive summers. Trees growing with Cistus as understory showed significant lower leaf water potential, leaf gas exchange parameters, leaf nitrogen content and chlorophyll photochemical efficiency than trees growing without shrub competence. However, the presence of the legume Retama did not affect significantly the physiological state of Q. ilex. Thus, we conclude that the presence of a shrubby understory has the potential to modify the functioning of scattered trees, but these effects are species-specific.     

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.     

Contrasting physiological responses of two co-occurring eucalypts to seasonal drought at restored bauxite mine sites

This study describes the physiological response of two co-occurring tree species (Eucalyptus marginata and Corymbia calophylla) to seasonal drought at low- and high-quality restored bauxite mine sites in south-western Australia. Seasonal changes in photosynthesis (A), stomatal conductance (g(s)), leaf water potential (ψ), leaf osmotic potential (ψ), leaf relative water content (RWC) and pressure-volume analysis were captured over an 18-month field study to (i) determine the nature and severity of physiological stress in relation to site quality and (ii) identify any physiological differences between the two species. Root system restriction at the low-quality site reduced maximum rates of gas exchange (g(s) and A) and increased water stress (midday ψ and daily RWC) in both species during drought. Both species showed high stomatal sensitivity during drought; however, E. marginata demonstrated a higher dehydration tolerance where ψ and RWC fell to -3.2 MPa and 73% compared with -2.4 MPa and 80% for C. calophylla. Corymbia calophylla showed lower g(s) and higher ψ and RWC during drought, indicating higher drought tolerance. Pressure-volume curves showed that cell-wall elasticity of E. marginata leaves increased in response to drought, while C. calophylla leaves showed lower osmotic potential at zero turgor in summer than in winter, indicating osmotic adjustment. Both species are clearly able to tolerate seasonal drought at hostile sites; however, by C. calophylla closing stomata earlier in the drought cycle, maintaining a higher water status during drought and having the additional mechanism of osmotic adjustment, it may have a greater capacity to survive extended periods of drought.     

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.     

Responses of deciduous forest trees to severe drought in Central Europe

In 2003, Central Europe experienced the warmest summer on record combined with unusually low precipitation. We studied plant water relations and phenology in a 100-year- old mixed deciduous forest on a slope (no ground water table) near Basel using the Swiss Canopy Crane (SCC). The drought lasted from early June to mid September. We studied five deciduous tree species; half of the individuals were exposed to elevated CO(2) concentration ([CO(2)]) (530 ppm) using a free-air, atmospheric CO(2)-enrichment system. In late July, after the first eight weeks of drought, mean predawn leaf water potential about 30 m above ground was -0.9 MPa across all trees, dropping to a mean of -1.5 MPa in mid-August when the top 1 m of the soil profile had no plant accessible moisture. Mean stomatal conductance and rates of maximum net photosynthesis decreased considerably in mid-August across all species. However, daily peak values of sap flow remained surprisingly constant over the whole period in Quercus petraea (Matt.) Liebl., and decreased to only about half of the early summer maxima in Fagus sylvatica L. and Carpinus betulus L. (stomatal down- regulation of flux). Although we detected no differences in most parameters between CO(2)-treated and control trees, predawn leaf water potential tended to be less negative in trees exposed to elevated [CO(2)]. Leaf longevity was greater in 2003 compared with the previous years, but the seasonal increase in stem basal area reached only about 75% of that in previous years. Our data suggest that the investigated tree species, particularly Q. petraea, did not experience severe water stress. However, an increased frequency of such exceptionally dry summers may have a more serious impact than a single event and would give Q. petraea a competitive advantage in the long run.     

Disturbances in European beech water relation during an extreme drought

Context

In the context of a probable increase in intensity and frequency of extreme summer drought events, a better understanding of the key processes involved in water relations is needed to improve the theoretical foundations of predictive process-based models.

Aims

This paper aims to analyse how temperate deciduous trees cope with water shortage.

Methods

The exceptional summer drought of 2003 in Europe provided an opportunity to monitor stomatal conductance and twig water potential in European beech (Fagus sylvatica L.) at predawn and midday and to analyse variations with respect to leaf height within the canopy. By comparing our field measurements of twig water potential to values found in the literature, we confirmed the strong impact of soil water shortage on crown water relations.

Results

This paper shows that (1) the vertical gradient of stomatal conductance within the crown disappeared under extreme soil water depletion; (2) at maximum drought intensity, predawn twig water potential (ψ _pd) reached ?2.3 MPa at a height of 14 m in the crown and ?2.0 MPa at a height of 10 m. The significant differences in ψ _pd between the two measurement heights in the canopy may be due to night transpiration; (3) there was a close relationship between predawn twig water potential and relative extractable soil water; (4) as drought conditions intensified, there was a close relationship between canopy radiation interception and predawn water potential, as estimated daily from relative extractable soil water.     

Sap flow index as an indicator of plant water status

Night and especially predawn tree water status is an important indicator of drought stress in trees. Leaf water potential (LWP) is frequently used as a measure of tree water status and hence drought stress; however, there are difficulties associated with sampling foliage from tall trees and determining LWP automatically. The current study was undertaken to determine whether sap flow index (SFI), which can be automatically and continuously recorded even during very low flows, can be used to estimate drought stress in trees caused by dry air under non-limiting soil water conditions. We made simultaneous measurements of LWP, heat pulse velocity (HPV) and SFI on apple trees (Malus domestica Borkh.) in the semiarid climate of southern Ukraine over several growing seasons. Predawn values of LWP were highly correlated with SFI. Over the range of low sap flow rates occurring at nighttime, where other methods of measuring sap flow are not sensitive, the SFI method was linear and very sensitive. Additional information about tree water status was obtained by comparing nighttime and daytime values of SFI. The ratio of predawn SFI to midday SFI and the period between the two daily SFI maxima (the first SFI peak occurred in the morning and the second peak occurred in the evening on cloudless days) can be used to characterize internal plant water balance. Although the daily course of SFI was variable, specific patterns were identified that reflected particular stages in the development of plant drought stress. An "air-drought-stress curve" was used to characterize the development of water stress in trees subjected to air drought during the growing season.     

Influences of Thinning and Prescribed Fire on Water Relations of Jeffrey Pine I. Xylem and Soil Water Potentials

Abstract

Forest thinning accomplished with cut-to-length and whole-tree harvesting systems, and prescribed underburning were assessed for their impacts on water relations in eastern Sierra Nevada Jeffrey pine (Pinus Jeffreyi Grev. & Balf.) during a period of extended drought. Predawn and midday measurements of xylem water potential in dominant and codominant crown class trees more than a century old were made on six days spread over three growing seasons, accompanied by measurements of soil water potential completed between the predawn and midday sessions of each day. With the exception of a single predawn session, the only one of a total of 12 in which xylem water potentials did not differ among treatments, the potentials in trees of thinned stand subunits were 0.67 MPa higher on average during predawn sessions and 0.71 MPa higher during midday sessions than those in trees of the unthinned treatment. Differences between the cut-to-length and whole-tree treatments were marginal and uncommon, but when they occurred, potentials were higher in the former. Prescribed fire effects on xylem water potential were also uncommon, but when occurring generally indicated lower stress levels in the burned than in the unburned treatment. Soil water potentials largely coincided with those of xylem water, with higher potentials in either the cut-to-length or whole-tree treatments, and usually both, than in the unthinned treatment on each of the six days of measurement. Underburning influences on soil water were rare, but when evident, potentials were higher in the burned than in the unburned treatment by substantial margins. For a majority of the measurement sessions, xylem water potential was found to be negatively correlated with residual basal area but positively correlated with soil water potential. In turn, coarse fragments and organic matter in the soil profile intermittently influenced soil water potential, with the former a negative factor while the latter was positive. Overall, results of this study suggest that substantial ecophysiological advantages can be derived from density management in older, dry site forests, which at minimum are not compromised by subsequent implementation of controlled underburning.     

宝天曼三桠乌药对降雨减少后的生理生态响应

[目的]在宝天曼森林原位建立降水减少(截雨)样地后,样地内的三桠乌药出现了顶端枯死,本研究从水和碳的角度,探讨三桠乌药顶端枯死的原因。[方法]2013年4月在宝天曼锐齿栎林原位建立了3块截雨样地,在2014年生长季干旱时,采用压力室、低压液流系统和蒽酮硫酸法测定了三桠乌药的水力系统特征和非结构性碳等指标。[结果]显示:(1)三桠乌药在截雨处理后出现了顶端枯死;三桠乌药最大导管长度约为60 cm,栓塞50%时的水势(P50)为-1.43 MPa,其木质部栓塞脆弱性较大。在截雨处理1年后生长季较干旱时,净光合速率、凌晨水势、中午水势显著低于对照,中午栓塞显著高于对照,且水力安全边际为负值。(2)三桠乌药气孔导度、蒸腾速率、叶面积、导管直径、边材比导率、叶片比导率显著下降,Huber值、导管密度显著升高;而叶片、韧皮部、木质部3个器官的可溶性糖、淀粉、总非结构性碳与对照差异不显著。[结论]水力失衡是三桠乌药顶端枯死的主要原因,而蒸腾面积、水分输导系统等的变化表明三桠乌药在缺水环境下进行了一定的适应性调节,但这些调节不足以使三桠乌药在缺水情况下避免水力失衡而导致的顶端枯死。     

Site-specific water relations and stomatal response of Quercus ilex in a Mediterranean watershed

Intraspecific variations in the water relations and stomatal response of Quercus ilex L. were analyzed under field conditions by comparing trees at two locations within a Mediterranean watershed (l'Avic, Catalonia, NE Spain). Distinct environmental gradients exist between the two sites (referred to as ridge top at 975 m and valley bottom at 700 m) with greater soil depth for water storage, reduced radiation, reduced wind and higher water vapor pressure deficits at the valley bottom than at the ridge top. Osmotic adjustment and changes in tissue elasticiity did not significantly increase drought resistance in the trees studied. The leaf-to-air vapor pressure difference (Deltaw) threshold for inducing stomatal closure was higher at the ridge top (15.6 kPa MPa(-1) +/- 0.5 SE) than at the valley bottom (9.8 kPa MPa(-1) +/- 1.0 SE). However, increases in Deltaw beyond the threshold were followed by greater reductions in leaf conductance of trees at the ridge top than at the valley bottom. At both sites, maximum leaf conductance was related to predawn shoot water potential which, in turn, was related to watershed stream flow. The effects of water deficits during the dry summer of 1989 were more severe in trees at the valley bottom than at the ridge top. During periods of high evaporative demand, site-specific differences in the control of water loss led to more conservative water use by trees at the ridge top and, thus, to even greater drought avoidance (higher predawn water potentials) in late summer.     

Water relations and gas exchange of Acer saccharum seedlings in contrasting natural light and water regimes

Field measurements were made of leaf photosynthesis (A), stomatal conductance (g) and leaf water relations for sugar maple (Acer saccharum Marsh.) seedlings growing in a forest understory, small gap or large clearing habitat in southwestern Wisconsin, USA. Predawn water status, leaf gas exchange and plasticity in field and laboratory water relations characteristics were compared among contrasting light environments in a wet year (1987) and a dry year (1988) to evaluate possible interactions between light and water availability in these habitats. Leaf water potentials (Psi(leaf)) at predawn and midday were lower for clearing than gap or understory seedlings. Acclimation of tissue osmotic potentials to light environment was observed among habitats but did not occur within any of the habitats in response to prolonged drought. During a summer drought in 1988, decreases in daily maximum g (g(max)) and maximum A (A(max)) in clearing seedlings were correlated with predawn Psi(leaf), which reached a seasonal minimum of -2.0 MPa. Under well-watered conditions, diurnal fluctuations in Psi(leaf) of up to 2.0 MPa in clearing seedlings occurred along with large midday depressions of A and g. In a wet year, strong stomatal responses to leaf-to-air vapor pressure difference (VPD) in sunny habitats were observed over nine diurnal courses of gas exchange measurements on seedlings in a gap and a clearing. Increasing stomatal limitations to photosynthesis appeared to be responsible for the reduction in A at high VPD for clearing seedlings. In understory seedlings, however, low water-use efficiency and development of leaf water deficits in sunflecks was related to reduced stomatal limitations to photosynthesis relative to seedlings in sunny habitats. Predawn Psi(leaf) and VPD appear to be important factors limiting carbon assimilation in sugar maple seedlings in light-saturating irradiances, primarily through stomatal closure. The overall results are consistent with the idea that sugar maple seedlings exhibit "conservative" water use patterns and have low drought tolerance. Leaf water relations and patterns of water use should be considered in studies of acclimation and species photosynthetic performance in contrasting light environments.     

Relationship of water status to vegetative growth and leaf gas exchange of peach (Prunus persica) trees on different rootstocks

We investigated relationships between tree water status, vegetative growth and leaf gas exchange of peach trees growing on different rootstocks under field conditions. Tree water status was manipulated by partially covering (0, approximately 30 and approximately 60%) the tree canopies on individual days and then evaluating the effects of tree water status on vegetative growth and leaf gas exchange. Early morning stem water potentials were approximately -0.4 MPa for trees in all treatments, but mean midday values ranged from -1.1 to -1.7 MPa depending on rootstock and canopy coverage treatment. Relative shoot extension growth rate, leaf conductance, transpiration rate and net CO2 exchange rate differed significantly among trees in the different rootstocks and canopy coverage treatments. Shoot extension growth rate, leaf conductance, leaf transpiration rate and leaf net CO2 exchange rate were linearly correlated with midday stem water potential. These relationships were independent of the rootstock and canopy coverage treatments, indicating that tree water relations are probably directly involved in the mechanism that imparts vegetative growth control by selected peach rootstocks.     

Osmotic regulation in leaves and roots of olive trees during a water deficit and rewatering

We evaluated the osmotic adjustment capacity of leaves and roots of young olive (Olea europaea L.) trees during a period of water deficit and subsequent rewatering. The trials were carried out in Basilicata (40 degrees 24' N, 16 degrees 48' E) on 2-year-old self-rooted olive plants (cv. 'Coratina'). Plants were subjected to one of four drought treatments. After 13 days of drought, plants reached mean predawn leaf water potentials of -0.45 +/- 0.015 MPa (control), -1.65 +/- 0.021 (low stress), -3.25 +/- 0.035 (medium stress) and -5.35 +/- 0.027 MPa (high stress). Total osmotic adjustment increased with increasing severity of drought stress. Trees in the high stress treatment showed total osmotic adjustments ranging between 2.4 MPa at 0500 h and 3.8 MPa at 1800 h on the last day of the drought period. Osmotic adjustment allowed the leaves to reach leaf water potentials of about -7.0 MPa. Active osmotic adjustment at predawn decreased during the rewatering period in both leaves and roots. Stomatal conductance and net photosynthetic rate declined with increasing drought stress. Osmotic adjustment in olive trees was associated with active and passive osmotic regulation of drought tolerance, providing an important mechanism for avoiding water loss.     

Mechanisms of drought response in Thuja occidentalis L. II. Post-conditioning water stress and stress relief

We examined the extent of osmotic adjustment and the changes in relative water content (RWC) and transpiration rate (i.e., relative stomatal function) that occur in water-deficit-conditioned 6-year-old Thuja occidentalis L. (eastern white cedar) trees in response to a severe drought. Trees conditioned by successive cycles of mild or moderate nonlethal water stress (conditioning) and nonconditioned trees were exposed to drought (i.e., -2.0 MPa predawn water potential) to determine if water deficit conditioning enhanced tolerance to further drought stress. Following drought, all trees were well watered for 11 days to evaluate how quickly osmotic potential, RWC and transpiration rate returned to preconditioning values. Both nonconditioned trees and mildly conditioned trees exhibited similar responses to drought, whereas moderately conditioned trees maintained higher water potentials and transpiration rates were 38% lower. Both conditioned and nonconditioned trees exhibited a similar degree of osmotic adjustment (-0.39 MPa) in response to drought relative to the well-watered control trees. The well-watered control trees, nonconditioned trees and mildly conditioned trees had similar leaf RWCs that were about 3% lower than those of the moderately conditioned trees. Following the 11-day stress relief, there were no significant differences in osmotic potential between the well-watered control trees and any of the drought-treated trees. Daily transpiration rates and water potential integrals (WPI) of all drought-treated trees approached those of the well-watered control trees during the stress relief period. However, the relationship between cumulative transpiration and WPI showed that previous exposure to drought stress reduced transpiration rates. Leaf RWC of the moderately conditioned trees remained slightly higher than that of the nonconditioned and mildly conditioned trees.     

Photosynthesis and water relations of the floodplain tree, boxelder (Acer negundo L.)

During summer, gas exchange and water relations were measured in mature boxelder (Acer negundo L.) trees growing on a floodplain in central Indiana, USA. A shallow (< 1.25-m deep) water table and repeated flooding kept the soil water potential above -0.5 MPa at all times. Net photosynthesis and stomatal conductance were influenced primarily by light and, to a lesser extent, by leaf temperature, but showed no relationships with leaf-to-air water vapor gradient or leaf water potential. Throughout the summer, there was no midday stomatal closure on any measurement day, and leaf water potential at dawn and minimum daily leaf water potential remained above -0.4 and -1.4 MPa, respectively. Nevertheless, there was a seasonal decline in leaf osmotic potentials at saturation and turgor-loss point. Seasonal changes in maximum daily net photosynthesis and stomatal conductance, minimum daily leaf water potential and soil-to-leaf hydraulic conductance were not related to seasonal changes in soil water potential, air or soil temperature, or water table depth. Seasonal responses of net photosynthesis to intercellular CO(2) indicated that net photosynthesis was controlled primarily by nonstomatal factors. High soil water and a shallow water table may have kept soil-to-leaf hydraulic conductance large (5-9 mmol m(-1) s(-1) MPa(-1)) throughout the summer, permitting the trees to keep their stomata open, yet maintain leaf turgor and high net photosynthesis during the hot, low-humidity afternoons. This could also account for the dominance of nonstomatal influences on net photosynthesis.     

Linking morphological and ecophysiological leaf traits to canopy dieback in Persian oak trees from central Zagros

Intraspecific variability in morphological and ecophysiological leaf traits might be theorized to be present in declining populations,since they seem to be exposed to stress and plasticity could be advantageous.Here we focused on declining Persian oaks(Quercus brantii Lindl.var.persica(Jaub and Spach)Zohary)in the Zagros Mountains of western Iran,representing the most important tree species of this region.We selected trees with contrasting crown dieback,from healthy to severely defoliated,to investigate the relationships between canopy dieback and leaf morphology,water content and pigments.We also measured esterase and peroxidase,as enzymatic antioxidants and indicators of contrasting genotypes.Trees showing moderate to severe defoliation showed higher leaf mass area(LMA),reduced relative water content(RWC),and lower stomatal density(SD).Increasing LMA indicates a more sclerophyllic structure,according to drier conditions.We did not find significant differences in leaf pigments(chlorophyll a and b,and carotenoids)among crown dieback classes,suggesting that Persian oak trees are able to maintain accurate photochemical efficiency,while reduced RWC and SD suggest hydraulic limitations.Our results do not provide a consistent pattern as regards enzymatic antioxidant defense in Persian oak.Morphological leaf traits would be important drivers of future adaptive evolution in Persian oak,leading to smaller and thicker leaves,which have fitness benefits in dry environments.Nonetheless,drought responses may be critically affecting carbon uptake,as photosynthetic compounds are less effectively used in leaves with higher sclerophylly.     

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.