Seasonal changes in apparent hydraulic conductance and their implications for water use of European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) and sessile oak [<Emphasis Type="Italic">Quercus petraea</Emphasis> (Matt.) Liebl] in South Europe

The water status of Fagus sylvatica L. and Quercus petraea (Matt) Liebl. was analysed during a cycle of progressive natural drought in southern Europe. Predawn (Ψ_pd) and midday water potential were measured in transpiring (Ψ_leaf) and non-transpiring leaves (Ψ_xyl). Furthermore, photosynthesis (A), stomatal conductance to water vapour (g_s) and sap flow (F_d) were recorded on the same dates. Apparent leaf specific hydraulic conductance in the soil–plant–air continuum (K_h) and whole tree hydraulic conductance (K_hsf) were calculated by using the simple analogy of the Ohm’s law. K_h was estimated at different points in the pathway as the ratio between transpiration (E) in the uppermost canopy leaves at midday and the gradient of water potential in the different compartments of the continuum soil–roots–stem–branches–leaves. There was a progressive decrease in water potential measured on non-transpiring leaves at the base of tree crown in both species (Ψ^l_xyl) from the beginning of the growing season to the end of summer. A similar decrease was shown in shoot water potential (Ψ^u_xyl) at the uppermost canopy. Predawn water potential (Ψ_pd) was high in both species until late July (28 July); afterwards, a significant decrease was registered in F. sylvatica and Q. petraea with minimum values of −0.81±0.03 and −0.75±0.06 MPa, respectively, by 15 September. In both species, leaf specific hydraulic conductance in the overall continuum soil–plant–air (K_h) decreased progressively as water stress increases. Minimum values of K_h and K_hsf were recorded when Ψ_pd was lower. However, Q. petraea showed higher K_h than F. sylvatica for the same Ψ_pd. The decrease in K_h with water stress was mainly linked to its fall from the soil to the lowermost canopy (K_srs). Nevertheless, a significant resistance in the petiole–leaf lamina (K_pl) was also recorded because significant differences in all dates were found on Ψ between transpiring and non-transpiring leaves from the same shoot. The decline in K_h was followed by an increase in stomatal control of daily water losses through the decrease of stomatal conductance to water vapour (g_s) during the day. It promoted a seasonal increase in the stomatal limitation to carbon dioxide uptake for photosynthesis (A). These facts were more relevant in F. sylvatica, which had concurrently a higher decline in water use at the tree level than Q. petraea. The results showed a strong coupling in F. sylvatica and Q. petraea between processes at leaf and tree level. It may be hypothesised a role of specific hydraulic conductance not only in the regulation of water losses by transpiration but also of carbon uptake.     

Water use patterns of four co-occurring chaparral shrubs

Summary Mixed stands of chaparral in California usually contain several species of shrubs growing close to each other so that aerial branches and subterranean roots overlap. There is some evidence that roots are stratified relative to depth. It may be that root stratification promotes sharing of soil moisture resources. We examined this possibility by comparing seasonal water use patterns in a mixed stand of chaparral dominated by four species of shrubs: Quercus durata, Heteromeles arbutifolia, Adenostoma fasciculatum, and Rhamnus californica. We used a neutron probe and soil phychrometers to follow seasonal depletion and recharging of soil moisture and compared these patterns to seasonal patterns of predawn water potentials, diurnal leaf conductances, and diurnal leaf water potentials. Our results indicated that 1) Quercus was deeply rooted, having high water potentials and high leaf conductances throughout the summer drought period, 2) Heteromeles/Adenostoma were intermediate in rooting depth, water potentials, and leaf conductances, and 3) Rhamnus was shallow rooted, having the lowest water potentials and leaf conductances. During the peak of the drought, predawn water potentials for Quercus corresponded to soil water potentials at or below a depth of 2 m, predawn water potentials of Heteromeles/ Adenostoma corresponded to a depth of 0.75 m, and predawn water potentials of Rhamnus corresponded to a depth of 0.5 m. This study supports the concept that co-occurring shrubs of chaparral in California utilize a different base of soil moisture resources.     

Comparative physiology of burned and unburned Rhus laurina after chaparral wildfire

Summary Laurel Sumac (Rhus laurina) is a dominant member of the coastal chaparral community of southern California that survives periodic burning by wildfires by resprouting from a lignotuber (root crown). We investigated the physiological basis for resprouting by comparing shoot elongation, leaf nitrogen content, tissue water status, leaf conductance to water vapor diffusion, and photosynthetic rates of post-fire R. laurina to those of adjacent unburned shrubs. Resprouts had higher rates of shoot elongation, leaf conductance, and photosynthesis than mature, unburned shrubs. Leaf nitrogen contents were elevated in burned shrubs even though their leaves developed interveinal chlorosis. A comparison of soil water potential to predawn water potential indicated that roots of R. laurina remain active below 2 m during the first summer drought after wildfire. Our results support the hypothesis that lignotubers not only contain dormant buds that develop into aerial shoots after wildfire but they also supply nutrient resources that enhance shoot elongation. Because R. laurina is relatively sensitive to drought, yet very successful in its rapid recovery after fire, maintaining an active root system after shoot removal may be the primary function of the massive lignotuber formed by this species.     

Gas exchange and photosynthesis of Eucalyptus camaldulensis seedlings inoculated with different ectomycorrhizal symbionts

Eucalyptus camaldulensis Dehnh. seedlings inoculated with Pisolithus tinctorius (Pers.) Coker & Couch and Thelephora terrestris Ehrl. per Fr. were grown in well watered soil (_s –0.03 MPa) or subjected to a long-term soil water stress of up to –1.0 MPa over 13-week period in a glasshouse. After 13 weeks, all seedling containers were watered to field capacity and then water was withheld from the E. camaldulensis seedlings to induce a short-term drought. Diurnal measurements of seedling photosynthesis rate (A), leaf stomatal conductance (g) and leaf water potential (_p) were completed before, during, and after the short term drought. Although they were growing in an equal soil volume, photosynthesis rate (A), leaf stomatal conductance and leaf water potential (_p) of larger seedlings with P. tinctorius ectomycorrhizae were similar to those of smaller seedlings colonized with T. terrestris during the short-term drought period. Seedlings inoculated with Pisolithus tinctorius maintained higher photosynthesis rates over the course of the short-term drought. Thus, P. tinctorius ectomycorrhizae appear to be more efficient than those of T. terrestris in assisting seedlings to maintain gas exchange and photosynthesis under limited soil moisture conditions.     

Bundle‐sheath cell regulation of xylem‐mesophyll water transport via aquaporins under drought stress: a target of xylem‐borne ABA?

The hydraulic conductivity of the leaf vascular system (K_leaf) is dynamic and decreases rapidly under drought stress, possibly in response to the stress phytohormone ABA, which increases sharply in the xylem sap (ABA_xyl) during periods of drought. Vascular bundle‐sheath cells (BSCs; a layer of parenchymatous cells tightly enwrapping the entire leaf vasculature) have been hypothesized to control K_leaf via the specific activity of BSC aquaporins (AQPs). We examined this hypothesis and provide evidence for drought‐induced ABA_xyl diminishing BSC osmotic water permeability (P_f) via downregulated activity of their AQPs. ABA fed to the leaf via the xylem (petiole) both decreased K_leaf and led to stomatal closure, replicating the effect of drought. In contrast, smearing ABA on the leaf blade, while also closing stomata, did not decrease K_leaf within 2–3 h of application, demonstrating that K_leaf does not depend entirely on stomatal closure. GFP‐labeled BSCs showed decreased P_f in response to ‘drought’ and ABA treatment, and a reversible decrease with HgCl₂ (an AQP blocker). These P_f responses, absent in mesophyll cells, suggest stress‐regulated AQP activity specific to BSCs, and imply a role for these cells in decreasing K_leaf via a reduction in P_f. Our results support the above hypothesis and highlight the BSCs as hitherto overlooked vasculature sensor compartments, extending throughout the leaf and functioning as ‘stress‐regulated valves’ converting vasculature chemical signals (possibly ABA_xyl) into leaf hydraulic signals.     

Leaf water characteristics and drought acclimation in sunflower genotypes

The responses of leaf water parameters to drought were examined using three sunflower (Helianthus annuus L.) genotypes. Osmotic potential at full water saturation (π¹⁰⁰), apoplastic water fraction (AWF) and bulk elastic modulus (BEM) were determined by pressure-volume curve analysis on well watered or on water-stressed plants (−1.0 MPa Ψ₁ < −1.5 MPa) previously drought-pretreated or not. The drought-pretreated plants were subjected to a 7-day drought period (predawn leaf water potential reached −0.9 MPa) followed by 8 days of rewatering. In well watered plants, all genotypes in response to drought acclimation displayed a significantly decreased π¹⁰⁰ associated with a decrease in the leaf water potential at the turgor-loss point (decrease in Ψ_tlp was between 0.15 and 0.21 MPa, depending on the genotype). In two genotypes, drought acclimation affected the partitioning of water between the apoplastic and symplastic fractions without any effect on the total amount of water in the leaves. As a third genotype displayed no modification of AWF and BEM after drought acclimation, the decreased π¹⁰⁰ was only due to the net accumulation of solutes and was consistent with the adjustment of the photochemical efficiency observed previously in this genotype in response to drought acclimation. In water-stressed plants, the osmotic adjustment (OA) can increase further beyond that observed in response to the drought pretreatment. However, the maintenance of photosynthetic rate and stomatal conductance at low leaf water potentials not only depends on the extent of osmotic adjustment, but also on the interaction between OA and AWF or BEM. Adaptative responses of leaf water parameters to drought are thus quite contrasted in sunflower genotypes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of flooding and drought on stomatal activity, transpiration, photosynthesis, water potential and water channel activity in strawberry stolons and leaves

Transpiration, xylem water potential and water channel activity were studied in developing stolons and leaves of strawberry (Fragaria × ananassa Duch.) subjected to drought or flooding, together with morphological studies of their stomata and other surface structures. Stolons had 0.12 stomata mm^–2 and a transpiration rate of 0.6 mmol H₂O m^–2 s^–1, while the leaves had 300 stomata mm^–2 and a transpiration rate of 5.6 mmol H₂O m^–2 s^–1. Midday water potentials of stolons were always less negative than in leaves enabling nutrient ion and water transport via or to the strawberry stolons. Drought stress, but not flooding, decreased stolon and leaf water potential from –0.7 to –1 MPa and from –1 to –2 MPa, respectively, with a concomitant reduction in stomatal conductance from 75 to 30 mmol H₂O m^–2 s^–1. However, leaf water potentials remained unchanged after flooding. Similarly, membrane vesicles derived from stolons of flooded strawberry plants showed no change in water channel activity. In these stolons, turgor may be preserved by maintaining root pressure, an electrochemical and ion gradient and xylem differentiation, assuming water channels remain open. By contrast, water channel activity was reduced in stolons of drought stressed strawberry plants. In every case, the effect of flooding on water relations of strawberry stolons and leaves was less pronounced than that of drought which cannot be explained by increased ABA. Stomatal closure under drought could be attributed to increased delivery of ABA from roots to the leaves. However, stomata closed more rapidly in leaves of flooded strawberry despite ABA delivery from the roots in the xylem to the leaves being strongly depressed. This stomatal closure under flooding may be due to release of stress ethylene. In the relative absence of stomata from the stolons, cellular (apoplastic) water transport in strawberry stolons was primarily driven by water channel activity with a gradient from the tip of the stolon to the base, concomitant with xylem differentiation and decreased water transport potential from the stolon tip to its base. Reduced water potential in the stolons under drought are discussed with respect to reduced putative water channel activity.     

Photosynthesis and stomatal conductance of potato leaves—effects of leaf age,irradiance, and leaf water potential

Potatoes (Solanum tuberosum L., cv. Bintje) were grown in a naturally lit glasshouse. Laboratory measurements on leaves at three insertion levels showed a decline with leaf age in photosynthetic capacity and in stomatal conductance at near saturating irradiance. Conductance declined somewhat more with age than photosynthesis, resulting in a smaller internal CO₂ concentration in older relative to younger leaves. Leaves with different insertion number behaved similarly. The changes in photosynthesis rate and in nitrogen content with leaf age were closely correlated. When PAR exceeded circa 100 W m^–2 the rate of photosynthesis and stomatal conductance changed proportionally as indicated by a constant internal CO₂ concentration. The photosynthesis-irradiance data were fitted to an asymptotic exponential model. The parameters of the model are AMAX, the rate of photosynthesis at infinite irradiance, and EFF, the slope at low light levels. AMAX declined strongly with leaf age, as did EFF, but to a smaller extent. During drought stress photosynthetic capacity declined directly with decreasing water potential (range –0.6 to –1.1 MPa). Initially, stomatal conductance declined faster than photosynthetic capacity.Abbreviations LNx leaf number x, counted in acropetal direction - DAP days after planting - DALA days after leaf appearance - C_i CO₂ concentration in the leaf - C_a CO₂ concentration in ambient air - LWP leaf water potential - OP osmotic potential - PAR photosynthetically active radiation     

Does engineering abscisic acid biosynthesis in Nicotiana plumbaginifolia modify stomatal response to drought?

The consequences of manipulating abscisic acid (ABA) biosynthesis rates on stomatal response to drought were analysed in wild‐type, a full‐deficient mutant and four under‐producing transgenic lines of N. plumbaginifolia. The roles of ABA, xylem sap pH and leaf water potential were investigated under four experimental conditions: feeding detached leaves with varying ABA concentration; injecting exogenous ABA into well‐watered plants; and withholding irrigation on pot‐grown plants, either intact or grafted onto tobacco. Changes in ABA synthesis abilities among lines did not affect stomatal sensitivity to ABA concentration in the leaf xylem sap ([ABA]_xyl), as evidenced with exogenous ABA supplies and natural increases of [ABA]_xyl in grafted plants subjected to drought. The ABA‐deficient mutant, which is uncultivable under normal evaporative demand, was grafted onto tobacco stock and then presented the same stomatal response to [ABA]_xyl as wild‐type and other lines. This reinforces the dominant role of ABA in controlling stomatal response to drought in N. plumbaginifolia whereas roles of leaf water potential and xylem sap pH were excluded under all studied conditions. However, when plants were submitted to soil drying onto their own roots, stomatal response to [ABA]_xyl slightly differed among lines. It is suggested, consistently with all the results, that an additional root signal of soil drying modulates stomatal response to [ABA]_xyl.     

Daily and seasonal variation in water relations of macchia shrubs and trees in France (Montpellier) and Turkey (Antalya)

Based upon two different research studies in the mediterranean regions of France and Turkey, drought resistance strategies were investigated in a broad group of species. The diurnal and seasonal patterns of the water relations of different lifeforms from the thermo-mediterranean to submediterranean lifezones were compared. Three sites near Montpellier, in Southern France, and five sites near Antalya, Turkey were used for this comparison. Xylem pressure potential and relative stomatal aperture were the key water relations parameters collected in France while these parameters as well as osmotic potential and leaf conductance were studied in Turkey.From the 26 different study species investigated in France, 7 distinct types of stomatal control were observed, with the deciduous lifeforms showing the least control, the sclerophyllous and coniferous evergreens the greatest control and the malacophyllous shrublets intermediate levels of control. Predawn water potential values provided a means of classifying species according to their temporal and spatial utilization of site water reserves. The comparison of turgor potentials (difference between water and osmotic potentials) gave an insight into leaf adaptations to site moisture. Species with high predawn water potentials generally maintain positive turgor even at midday during the summer, whereas species with low predawn values were frequently at zero turgor even at predawn. Phlomis grandiflora was the most extreme species with mid-summer predawns and midday water potentials of –6 MPa and osmotic potentials never more negative than –2.4 MPa.     

Is stomatal conductance in a tomato crop controlled by soil or atmosphere?

Summary The effects of soil water deficits and air vapour pressure deficits on stomatal conductance of tomato leaves were analysed separately under field conditions in central Portugal. Three conditions were created: low soil and air humidity (A), high soil and air humidity (B) and low soil but high air humidity (C). The results show that the effect of air vapour pressure deficit on stomatal behaviour is more important than the effect of soil water deficit when the predawn leaf water potential is above –0.4 MPa.     

Influence of Seasonal Variations in Soil Water Availability on Gas Exchange of Tropical Canopy Trees

Seasonal variations in environmental conditions influence the functioning of the whole ecosystem of tropical rain forests, but as yet little is known about how such variations directly influence the leaf gas exchange and transpiration of individual canopy tree species. We examined the influence of seasonal variations in relative extractable water in the upper soil layers on predawn leaf water potential, saturated net photosynthesis, leaf dark respiration, stomatal conductance, and tree transpiration of 13 tropical rain forest canopy trees (eight species) over 2 yr in French Guiana. The canopies were accessed by climbing ropes attached to the trees and to a tower. Our results indicate that a small proportion of the studied trees were unaffected by soil water depletion during seasonal dry periods, probably thanks to efficient deep root systems. The trees showing decreased tree water status (i.e., predawn leaf water potential) displayed a wide range of leaf gas exchange responses. Some trees strongly regulated photosynthesis and transpiration when relative extractable water decreased drastically. In contrast, other trees showed little variation, thus indicating good adaptation to soil drought conditions. These results have important applications to modeling approaches: indeed, precise evaluation and grouping of these response patterns are required before any tree‐based functional models can efficiently describe the response of tropical rain forest ecosystems to future changes in environmental conditions.     

Influence of cold soil and snowcover on photosynthesis and leaf conductance in two Rocky Mountain conifers

Summary The influence of cold soil and snowcover on photosynthesis and conductance of Picea engelmannii and Pinus contorta was investigated early in the growing season in the Medicine Bow Mountains, Wyoming, USA. Trees of both species growing in cold soil (<1°C) associated with snowpack had 25–40% lower leaf photosynthesis than trees in warm soils (>10°C). In cold soils leaf conductance of both species was lower, but more so in Pinus, leading to lower intercellular CO₂ concentrations and greater stomatal limitation of photosynthesis. Soil temperature had no effect on predawn and midday shoot water potentials of Pinus and Picea and lower photosynthesis and conductance did not appear to be a result of lower bulk shoot water potential. Predawn, as well as midday, water potentials of Pinus were consistently higher than Picea suggesting that Pinus may have deeper roots, although trenching experiments indicated young Picea trees have more extensive lateral root systems than similar sized Pinus trees. Young Picea trees (<2 m in height) in snowbanks were capable of utilizing warmer soil 4 m from their base. Under similar conditions Pinus in snowbanks had lower photosynthesis and conductance than controls and Pinus did not appear capable of utilizing warmer soils nearby. Under full sunlight, PPFD reflected from the snow surface was 400–1400 mol m^-2 s^-1 higher than from snow-free surfaces. This reflected light resulted in a 10%–20% increase in photosynthesis of Picea. The beneficial effect of reflected light was apparent whether or not photosynthesis was reduced by low soil temperatures.     

Drought acclimation of two deciduous tree species of different layers in a temperate forest canopy

Water-use strategies of Populus tremula and Tilia cordata, and the role of abscisic acid in these strategies, were analysed. P. tremula dominated in the overstorey and T. cordata in the lower layer of the tree canopy of the temperate deciduous forest canopy. Shoot water potential (), bulk-leaf abscisic acid concentration ([ABA]_leaf), abscisic acid concentration in xylem sap ([ABA]_xyl), and rate of stomatal closure following the supply of exogenous ABA (v) decreased acropetally through the whole tree canopy, and foliar water content per area (w), concentration of the leaf osmoticum (c), maximum leaf-specific hydraulic conductance of shoot (L), stomatal conductance (g_s), and the threshold dose per leaf area of the exogenous ABA (d_a) required to reduce stomatal conductance increased acropetally through the tree canopy (from the base of the foliage of T. cordata to the top of the foliage of P. tremula) in non-stressed trees. The threshold dose per leaf dry mass of the exogenous ABA (d_w) required to reduce stomatal conductance, was similar through the tree canopy. After a drought period (3 weeks), the , w, L, g_s, d_a and d_w had decreased, and c and v had increased in both species. Yet, the effect of the drought period was more pronounced on L, g_s, d_a, d_w and v in T. cordata, and on , w and c in P. tremula. It was concluded that the water use of the species of the lower canopy layer—T. cordata, is more conservative than that of the species of the overstorey, P. tremula. [ABA]_leaf had not been significantly changed in these trees, and [ABA]_xyl had increased during the drought period only in P. tremula. The relations between [ABA]_leaf, [ABA]_xyl and the stomatal conductance, the osmotic adjustment and the shoot hydraulic conductance are also discussed.     

Drought controls over conductance and assimilation of a Mediterranean evergreen ecosystem: scaling from leaf to canopy

Drought control over conductance and assimilation was assessed using eddy flux and meteorological data monitored during four summer periods from 1998 to 2001 above a closed canopy of the Mediterranean evergreen oak tree Quercus ilex. Additional discrete measurements of soil water content and predawn leaf water potential were used to characterize the severity of the drought. Canopy conductance was estimated through the big‐leaf approach of Penman–Monteith by inverting latent heat fluxes. The gross primary production ( GPP ) was estimated by adding ecosystem respiration to net ecosystem exchange. Ecosystem respiration was deduced from night flux when friction velocity ( u _*) was greater than 0.35 m s^?1. Empirical equations were identified that related maximal canopy conductance and daily ecosystem GPP to relative soil water content ( RWC) , the ratio of current soil water content to the field capacity, and to the predawn leaf water potential. Both variables showed a strong decline with soil RWC for values lower than 0.7. The sharpest decline was observed for GPP . The curves reached zero for RWC =0.41 and 0.45 for conductance and GPP , respectively. When the predawn leaf water potential was used as a surrogate for soil water potential, both variables showed a hyperbolic decline with decreasing water potential. These results were compared with already published literature values obtained at leaf level from the same tree species. Scaling up from the leaf to ecosystem highlighted the limitation of two big‐leaf representations: Penman–Monteith and Sellers' Π factor. Neither held completely for comparing leaf and canopy fluxes. Tower measurements integrate fluxes from foliage elements clumped at several levels of organization: branch, tree, and ecosystem. The Q. ilex canopy exhibited non‐random distribution of foliage, emphasizing the need to take into account a clumping index, the factor necessary to apply the Lambert–Beer law to natural forests. Our results showed that drought is an important determinant in water losses and CO₂ fluxes in water‐limited ecosystems. In spite of the limitations inherent to the big‐leaf representation of the canopy, the equations are useful for predicting the influence of environmental factors in Mediterranean woodlands and for interpreting ecosystem exchange measurements.     

CO2 alters water use,carbon gain,and yield for the dominant species in a natural grassland

Global atmospheric CO₂ is increasing at a rate of 1.5–2 ppm per year and is predicted to double by the end of the next century. Understanding how terrestrial ecosystems will respond in this changing environment is an important goal of current research. Here we present results from a field study of elevated CO₂ in a California annual grassland. Elevated CO₂ led to lower leaf-level stomatal conductance and transpiration (approximately 50%) and higher mid-day leaf water potentials (30–35%) in the most abundant species of the grassland, Avena barbata Brot. Higher CO₂ concentrations also resulted in greater midday photosynthetic rates (70% on average). The effects of CO₂ on stomatal conductance and leaf water potential decreased towards the end of the growing season, when Avena began to show signs of senescence. Water-use efficiency was approximately doubled in elevated CO₂, as estimated by instantaneous gas-exchange measurements and seasonal carbon isotope discrimination. Increases in CO₂ and photosynthesis resulted in more seeds per plant (30%) and taller and heavier plants (27% and 41%, respectively). Elevated CO₂ also reduced seed N concentrations (9%).     

Different responses to drought and freeze stress of Quercus ilex L. growing along a latitudinal gradient

The vulnerability to drought and freeze stress was measured in young plants of Quercus ilex L. growing in the field in two natural sites within the Italian distribution area of this species, i.e. Sicily (Southern Mediterranean Basin) and Venezia Giulia (Northeastern Italy), respectively. In particular, the resistance strategies adopted by Q. ilex to withstand the two stresses were estimated in terms of seasonal and/or diurnal changes in leaf conductance to water vapour (g_L), water potential (_L) and relative water content (RWC) as well as of xylem embolism in the stem and root hydraulic conductance (K_RL). Sicilian (SI) plants showed to reduce water loss by stomatal closure (g_L decreased) in summer, thus maintaining average RWCs at 88–90%. Moreover, SI plants showed considerable resistance to xylem cavitation in the stem (the loss of hydraulic conductance, PLC, was less than 12% throughout the year) and to maintain the hydraulic conductance of their roots (K_RL), constantly high even in summer. Plants growing in Venezia Giulia (VG plants), on the contrary, underwent leaf dehydration in the winter due to freeze stress so that RWC measured in April was still 78% on a diurnal basis. This was apparently due to consistent xylem embolism in the stem. In fact, PLC was as high as 40% between November and March. Only in the summer was PLC similar to that recorded in SI plants. Moreover, K_RL of VG plants decreased in November from about 1.5 to 0.8×10^–4 kg s^–1 m^–2 MPa^–1, i.e. about 50%, and in February K_RL dropped further to 0.4×10^–4 kg s^–1 m^–2 MPa^–1. On the basis of the above, we conclude that: (a) Q. ilex was more sensitive to freeze than to drought stress so that freeze stress can be considered as a factor limiting the distribution area of this species; (b) drought and freeze stress were faced by Q. ilex adopting two different resistance strategies, i.e. drought avoidance based on water saving in Sicily and freeze tolerance in Venezia Giulia.     

Paraheliotropic leaf movement in Siratro as a protective mechanism against drought-induced damage to primary photosynthetic reactions: damage by excessive light and heat

Damage to primary photosynthetic reactions by drought, excess light and heat in leaves of Macroptilium atropurpureum Dc. cv. Siratro was assessed by measurements of chlorophyll fluorescence emission kinetics at 77 K (-196°C). Paraheliotropic leaf movement protected waterstressed Siratro leaves from damage by excess light (photoinhibition), by heat, and by the interactive effects of excess light and high leaf temperatures. When the leaves were restrained to a horizontal position, photoinhibition occurred and the degree of photoinhibitory damage increased with the time of exposure to high levels of solar radiation. Severe inhibition was followed by leaf death, but leaves gradually recovered from moderate damage. This drought-induced photoinhibitory damage seemed more closely related to low leaf water potential than to low leaf conductance. Exposure to leaf temperatures above 42°C caused damage to the photosynthetic system even in the dark and leaves died at 48°C. Between 42 and 48°C the degree of heat damage increased with the time of exposure, but recovery from moderate heat damage occurred over several days. The threshold temperature for direct heat damage increased with the growth temperature regime, but was unaffected by water-stress history or by current leaf water status. No direct heat damage occurred below 42°C, but in water-stressed plants photoinhibition increased with increasing leaf temperature in the range 31–42°C and with increasing photon flux density up to full sunglight values. Thus, water stress evidently predisposes the photosynthetic system to photoinhibition and high leaf temperature exacerbates this photoinhibitory damage. It seems probable that, under the climatic conditions where Siratro occurs in nature, but in the absence of paraheliotropic leaf movement, photoinhibitory damage would occur more frequently during drought than would direct heat damage.Abbreviations and symbols PFD photon flux area density - PSI, PSII photosyntem I, II - F _M, F _O, F _V maximum, instantaneous, variable fluorescence emission - PLM paraheliotropic leaf movement; all data of parameter of variation are mean ± standard error     

Drought stress effects on three cultivars of Eragrostis curvula: photosynthesis and water relations

Water stress effects were studied on three cultivars ofEragrostis curvula. Leaf water potential, RWC, total plantleaf area, green dry weight mass percentage and CO₂ gas-exchangeweremeasured during the onset of stress and after recovery. After 3 days of waterstress, RWC of cv Tanganyika plants was around 30–40% of controls,while RWC of cvs Ermelo and Consol was around 50–60% of controls.However midday and predawn water potentials were lower in cvs Tanganyka andErmelo than in cv Consol. After re-watering, RWC and water potentials recoveredonly in Consol plants. A strong decrease of leaf area was recorded in cvsErmeloand Consol during water stress (about 91–94% less than the leafarea of controls). Photosynthesis decreased as a function of the degree ofwaterstress severity in all cultivars. Also, light saturated photosynthesis,CO₂ quantum yield and light at which saturated photosynthesisoccurred, were strongly reduced by water stress. Recovery of photosynthesis wasfound in cv Consol after five days re-watering. Cv Consol showed a betterconservation of water and higher resistance to water stress than the other twocvs.     

Transpiration and stomatal behaviour of Quercus ilex plants during the summer in a Mediterranean carbon dioxide spring

Variations in the water relations and stomatal response of Quercus ilex were analysed under field conditions by comparing trees at two locations in a Mediterranean environment during two consecutive summers (1993 and 1994). We used the heat-pulse velocity technique to estimate transpirational water use of trees during a 5 month period from June to November 1994. At the end of sap flow measurements, the trees were harvested, and the foliage and sapwood area measured. A distinct environmental gradient exists between the two sites with higher atmospheric CO₂ concentrations in the proximity of a natural CO₂ spring. Trees at the spring site have been growing for generations in elevated atmospheric CO₂ concentrations. At both sites, maximum leaf conductance was related to predawn shoot water potential. The effects of water deficits on water relations and whole-plant transpiration during the summer drought were severe. Leaf conductance and water potential recovered after major rainfall in September to predrought values. Sap flow, leaf conductance and predawn water potential decreased in parallel with increases in hydraulic resistance, reaching a minimum in mid-summer. These relationships are in agreement with the hypothesis of the stomatal control of transpiration to prevent desiccation damage but also to avoid ‘runaway embolism’. Trees at the CO₂ spring underwent less reduction in hydraulic resistance for a given value of predawn water potential. The decrease in leaf conductance caused by elevated CO₂ was limited and tended to be less at high than at low atmospheric vapour pressure deficit. Mean (and diurnal) sap flux were consistently higher in the control site trees than in the CO₂ spring trees. The degree of reduction in water use between the two sites varied among the summer periods. The control site trees had consistently higher sap flow at corresponding values of either sapwood cross-sectional area or foliage area. Larger trees displayed smaller differences than smaller trees, between the control and the CO₂ spring trees. A strong association between foliage area and sapwood cross-sectional area was found in both the control and the CO₂ spring trees, the latter supporting a smaller foliage area at the corresponding sapwood stem cross-sectional area. The specific leaf area (SLA) of the foliage was not influenced by site. The results are discussed in terms of the effects of elevated CO₂ on plant water use at the organ and whole-tree scale.