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.     

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.     

Diurnal changes in water conduction in loblolly pine (Pinus taeda) and Virginia pine (P. virginiana) during soil dehydration

We studied diurnal changes in water conduction during soil dehydration in 37-month-old seedlings of one Virginia pine (Pinus virginiana Mill.) and two loblolly pine (P. taeda L.) sources, one from North Carolina (NC) and the other from the "Lost Pines" areas of Texas (TX), in an environmentally controlled growth chamber. For seedlings of similar biomass, the TX source had higher values of transpiration, needle conductance, and plant hydraulic conductivity under well-watered conditions than the NC source. Under dry soil conditions, the TX source had lower values of water conduction than the NC source. The Virginia pine source responded similarly to the TX source under both well-watered and dry soil conditions. For all three pine sources, gradients between soil and needle water potentials were greatest when the seedlings were moderately stressed. The TX and Virginia pine sources had higher gradients and lower daytime needle water potentials under moderate stress conditions than the NC source. Predawn needle water potentials did not differ among the pine sources. We conclude that the TX and Virginia pine sources have decreased daytime needle water potentials and increased water potential gradients during the daytime under moderate stress conditions, but with no disruption of recovery at predawn. The greater rates of transpiration and water conduction by the TX source compared with the NC source under well-watered conditions suggest a means by which growth can be maximized prior to the onset of drought, thereby enhancing survival of loblolly pines in drought-prone environments.     

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.     

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.     

The response of Pinus sylvestris to drought: stomatal control of transpiration and hydraulic conductance

We investigated the impact of drought on the physiology of 41-year-old Scots pine (Pinus sylvestris L.) in central Scotland. Measurements were made of the seasonal course of transpiration, canopy stomatal conductance, needle water potential, xylem water content, soil-to-needle hydraulic resistance, and growth. Comparison was made between drought-treated plots and those receiving average precipitation. In response to drought, transpiration rate declined once volumetric water content (VWC) over the top 20 cm of soil reached a threshold value of 12%. Thereafter, transpiration was a near linear function of soil water content. As the soil water deficit developed, the hydraulic resistance between soil and needles increased by a factor of three as predawn needle water potential declined from -0.54 to -0.71 MPa. A small but significant increase in xylem embolism was detected in 1-year-old shoots. Stomatal control of transpiration prevented needle water potential from declining below -1.5 MPa. Basal area, and shoot and needle growth were significantly reduced in the drought treatment. In the year following the drought, canopy stomatal conductance and soil-to-needle hydraulic resistance recovered. Current-year needle extension recovered, but a significant reduction in basal area increment was evident one year after the drought. The results suggest that, in response to soil water deficit, mature Scots pine closes its stomata sufficiently to prevent the development of substantial xylem embolism. Reduced growth in the year after a severe soil water deficit is most likely to be the result of reduced assimilation in the year of the drought, rather than to any residual embolism carried over from one year to the next.     

Interspecific variation in leaf water use associated with drought tolerance in four emergent dipterocarp species of a tropical rain forest in Borneo

By use of tree-tower and canopy-crane systems we studied variations in the water use, including transpiration, stomatal conductance, and leaf water potential, of the uppermost sun-exposed canopy leaves of four emergent dipterocarp species in an aseasonal tropical rain forest in Sarawak, Malaysia. Midday depression in stomatal conductance and leaf water potential was observed in all the species studied. Interspecific differences were clearly observed in the maxima of transpiration rates and stomatal conductance and the minima of leaf water potential among the four dipterocarp species. These interspecific variations were closely related to wood density and to factors affecting ecological patterns of distribution. Specifically, Shorea parvifolia and S. smithiana, both of which have a relatively low wood density for Dipterocarpaceae and are found on clay-rich soil, had a high transpiration rate in the daytime but had a large midday depression and a low leaf water potential. In contrast, Dryobalanops aromatica, which has a high wood density and is found in sandy soil areas, consumed less water even during the daytime. Dipterocarpus pachyphyllus, which has a high wood density and is found on clay-rich soil, stood intermediate between Shorea and D. aromatica in leaf water use. The two Shorea species had higher mortality than the others during the severe drought associated with El Ni?o in 1998, so daily pattern of leaf water use in each dipterocarp species might be correlated with its susceptibility to unusual drought events.     

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.     

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.     

Xylem sap flow and drought stress of Fraxinus excelsior saplings

We report on diurnal and seasonal variations in sap flow rate and stem water potential of Fraxinus excelsior L. saplings growing at the edge of a Fraxino-Aceretum forest in western Germany. Because of shallow soil, the trees were subjected to drought in summer. When soil water availability was not limiting, sap flow rate was related to changes in solar radiation and vapor pressure deficit. Maximum transpiration rates per leaf area were 3.5-7.4 mmol m-2 s-1, and maximum daily totals were 1.7-3.3 kg m-2 day-1. Under drought conditions, stem water potential dropped to midday minima of -2.6 to -3.5 MPa and sap flow rate was strongly related to this parameter. After the drought period, reduced apparent (whole-plant) hydraulic conductance was observed, which was attributed to a continued reduction in stomatal conductance after the drought stress had ceased. A model was developed that linked sap flow rate directly to climatic variables and stem water potential. Good correlation between measured and simulated sap flow rates allowed the model to be used for data interpretation.     

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.     

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.     

Processes preventing nocturnal equilibration between leaf and soil water potential in tropical savanna woody species

The impact of nocturnal water loss and recharge of stem water storage on predawn disequilibrium between leaf (psiL) and soil (psiS) water potentials was studied in three dominant tropical savanna woody species in central Brazil (Cerrado). Sap flow continued throughout the night during the dry season and contributed from 13 to 28% of total daily transpiration. During the dry season, psiL was substantially less negative in covered transpiring leaves, throughout the day and night, than in exposed leaves. Before dawn, differences in psiL between covered and exposed leaves were about 0.4 MPa. When relationships between sap flow and psiL of exposed leaves were extrapolated to zero flow, the resulting values of psiL (a proxy of weighted mean soil water potential) in two of the species were similar to predawn values of covered leaves. Consistent with substantial nocturnal sap flow, stomatal conductance (gs) never dropped below 40 mmol m(-2) s(-1) at night, and in some cases, rose to as much as 100 mmol m(-2) s(-1) before the end of the dark period. Nocturnal gs decreased linearly with increasing air saturation deficit (D), but there were species-specific differences in the slopes of the relationships between nocturnal gs and D. Withdrawal and recharge of water from stem storage compartments were assessed by monitoring diel fluctuations of stem diameter with electronic dendrometers. Stem water storage compartments tended to recharge faster when nocturnal transpiration was reduced by covering the entire plant. Water potential of covered leaves did not stabilize in any of the plants before the end of the dark period, suggesting that, even in covered plants, water storage tissues were not fully rehydrated by dawn. Patterns of sap flow and expansion and contraction of stems reflected the dynamics of water movement during utilization and recharge of stem water storage tissues. This study showed that nighttime transpiration and recharge of internal water storage contribute to predawn disequilibrium in water potential between leaves and soil in neotropical savanna woody plants.     

Variations in transpiration rate and leaf cell turgor maintenance in saplings of deciduous broad-leaved tree species common in cool temperate forests in Japan

To clarify mechanisms underlying variation in transpiration rate among deciduous broad-leaved tree species, we measured diurnal changes in stomatal conductance (gs) and leaf water potential, and calculated the maximum transpiration rate (Emax), leaf-specific hydraulic conductance (K(s-l)) and difference between the soil water potential and the daily minimum leaf water potential (Psis - Psi(l,min)). Pressure-volume (P-V) measurements were made on leaves. Saplings of eight broad-leaved tree species that are common in Japanese cool temperate forests were studied. Maximum transpiration rate varied significantly among species. There was a statistically significant difference in Psis - Psi(l,min), but not in K(s-l). Species with large Emax also had large Psis - Psi(l,min) and gs. The results of the P-V analyses showed that species with a large Psis - Psi(l,min) maintained turgor even at low leaf water potentials. The similar daily minimum leaf pressure potentials (Psip) across all eight species indicate that Psip values below this minimum are critical. Based on these results, we suggest that the leaf cell capacity for turgor maintenance strongly affects Psis - Psi(l,min) and consequently Emax via stomatal regulation.     

Chronic cork oak decline and water status: new insights

Chronic decline and Sudden death are two syndromes of cork oak (Quercus suber) dieback. Mortality is associated with water stress, but underlying physiological mechanisms are poorly understood. Here, we investigated the physiological performance of declining trees during the summer drought. Leaf water potential, gas-exchange, fluorescence of photosystem II and leaf and root starch concentration were compared in healthy (asymptomatic) and declining trees. Low annual cork increment in declining trees indicated tree decline for several years. All trees showed similar water status in spring. In summer, declining trees showed lower predawn leaf water potential (?2.0 vs. ?0.8 MPa), but unexpectedly higher midday leaf water potential than healthy trees (?2.8 vs. ?3.3 MPa). The higher midday water potential was linked to by means of strongly reduced stomatal conductance and, consequently, transpiration. This study is pioneer showing that declining trees had high midday water potential. A tendency for lower sap flow driving force (the difference between predawn and midday water potential) in declining trees was also associated with reduced photosynthesis, suggesting that chronic dieback may be associated with low carbon uptake. However, starch in roots and leaves was very low and not correlated to the health status of trees. Declining trees showed lower water-use efficiency and non-photochemical quenching in summer, indicating less resistance to drought. Contrarily to chronic decline, one tree that underwent sudden death presented predawn leaf water potential below the cavitation threshold.     

Genotypic variation in drought response of silver birch (Betula pendula): leaf water status and carbon gain

To assess genotypic variation in drought response of silver birch (Betula pendula Roth), we studied the plasticity of 16 physiological traits in response to a 12-14-week summer drought imposed on four clones in two consecutive years. In a common garden experiment, 1-year-old clonal trees from regions with low (550 mm year(-1)) to high rainfall (1270 mm year(-1)) were grown in 45-l pots, and leaf gas exchange parameters, leaf water potentials, leaf osmotic potentials and leaf carbon isotope signatures were repeatedly measured. There were no clonal differences in leaf water potential, but stomatal conductance (gs), net photosynthesis at ambient carbon dioxide concentration, photosynthetic water-use efficiency, leaf carbon isotope composition (delta13C) and leaf osmotic potentials at saturation (Pi0) and at incipient plasmolysis (Pip) were markedly influenced by genotype, especially gs and osmotic adjustment. Genotypes of low-rainfall origin displayed larger osmotic adjustment than genotypes of high-rainfall origin, although their Pi0 and Pip values were similar or higher with ample water supply. Genotypes of low-rainfall origin had higher gs than genotypes of high-rainfall origin under both ample and limited water supply, indicating a higher water consumption that might increase competitiveness in drought-prone habitats. Although most parameters tested were significantly influenced by genotype and treatment, the genotype x treatment interactions were not significant. The genotypes differed in plasticity of the tested parameters and in their apparent adaptation to drought; however, among genotypes, physiological plasticity and drought adaptation were not related to each other. Reduction of gs was the first and most plastic response to drought in all genotypes, and allowed the maintenance of high predawn leaf water potentials during the drought. None of the clones exhibited non-stomatal limitation of photosynthesis. Leaf gs, photosynthetic capacity, magnitude of osmotic adjustment and delta13C were all markedly lower in 2000 than in 1999, indicating root limitation in the containers in the second year.     

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.     

Stomatal limitation to CO2 assimilation and down-regulation of photosynthesis in Quercus ilex resprouts in response to slowly imposed drought

Holm oak (Quercus ilex L.) is native to hot, dry Mediterranean forests where limited water availability often reduces photosynthesis in many species, and forest fires are frequent. Holm oaks resprout after a disturbance, with improved photosynthetic activity and water relations compared with unburned plants. To better understand the role of water availability in this improvement, watering was withheld from container-grown plants, either intact (controls) or resprouts after excision of the shoot, to gradually obtain a wide range of soil water availabilities. At high water availability, gas exchange rates did not differ between controls and resprouts. At moderate soil dryness, net photosynthesis of control plants decreased as a result of increased stomatal limitation, whereas gas exchange rates of resprouts, which had higher midday and predawn leaf water potentials, were unchanged. Under severe drought, resprouts showed a less marked decline in gas exchange than controls and maintained photosystem II integrity, as indicated by chlorophyll fluorescence measurements. Photosynthesis was down-regulated in both plant types in response to reduced CO2 availability caused by high stomatal limitation. Lower non-stomatal limitations in resprouts than in control plants, as evidenced by higher carboxylation velocity and the capacity for ribulose-1,5-bisphosphate regeneration, conferred greater drought resistance under external constraints similar to summer conditions at midday.     

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.