Short-term effects of potassium fertilization on the hydraulic conductance of Laurus nobilis L

This study reports experimental evidence on the effect of short-term potassium fertilization on potassium uptake, tissue concentration and hydraulic conductance of pot-grown laurel plants. Potassium uptake and loading into the xylem of laurel seedlings increased within 24 h after fertilization. Potassium was not accumulated in roots and leaves, but the [K(+)] of xylem sap was 80% higher in fertilized plants (+K) than in potassium-starved plants (-K), as a likely result of recirculation between xylem and phloem. Increased xylem sap [K(+)] resulted in a 45% increase in transpiration rate, a 30% increase in plant hydraulic conductance (K(plant)) and a 120% increase in leaf-specific conductivity of the shoot (k(shoot)). We suggest that this increase was due to ion-mediated up-regulation of xylem hydraulics, possibly caused by the interaction of potassium ions with the pectic matrix of intervessel pits. The enhancement of hydraulic conductance following short-term potassium fertilization is a phenomenon that can be of advantage to plants for maintaining cell turgor, stomatal aperture and gas exchange rates under moderate drought stress. Our data provide additional support for the important role of potassium nutrition in agriculture and forestry.     

Co-optimal distribution of leaf nitrogen and hydraulic conductance in plant canopies

Leaf properties vary significantly within plant canopies, due to the strong gradient in light availability through the canopy, and the need for plants to use resources efficiently. At high light, photosynthesis is maximized when leaves have a high nitrogen content and water supply, whereas at low light leaves have a lower requirement for both nitrogen and water. Studies of the distribution of leaf nitrogen (N) within canopies have shown that, if water supply is ignored, the optimal distribution is that where N is proportional to light, but that the gradient of N in real canopies is shallower than the optimal distribution. We extend this work by considering the optimal co-allocation of nitrogen and water supply within plant canopies. We developed a simple 'toy' two-leaf canopy model and optimized the distribution of N and hydraulic conductance (K) between the two leaves. We asked whether hydraulic constraints to water supply can explain shallow N gradients in canopies. We found that the optimal N distribution within plant canopies is proportional to the light distribution only if hydraulic conductance, K, is also optimally distributed. The optimal distribution of K is that where K and N are both proportional to incident light, such that optimal K is highest to the upper canopy. If the plant is constrained in its ability to construct higher K to sun-exposed leaves, the optimal N distribution does not follow the gradient in light within canopies, but instead follows a shallower gradient. We therefore hypothesize that measured deviations from the predicted optimal distribution of N could be explained by constraints on the distribution of K within canopies. Further empirical research is required on the extent to which plants can construct optimal K distributions, and whether shallow within-canopy N distributions can be explained by sub-optimal K distributions.     

A potential role for xylem-phloem interactions in the hydraulic architecture of trees: effects of phloem girdling on xylem hydraulic conductance

We investigated phloem-xylem interactions in Acer rubrum L. and Acer saccharum Marsh. Our experimental method allowed us to determine xylem conductance of an intact branch by measuring the flow rate of water supplied at two delivery pressures to the cut end of a small side branch. We found that removal of bark tissue (phloem girdling) upstream of the point at which deionized water was delivered to the branch resulted in a decrease (24% for A. rubrum and 15% for A. saccharum) in branch xylem hydraulic conductance. Declines in hydraulic conductance with girdling were accompanied by a decrease in the osmotic concentration of xylem sap. The decrease in xylem sap concentration following phloem girdling suggests that ion redistribution from the phloem was responsible for the observed decline in hydraulic conductance. When the same measurements were made on branches perfused with KCl solution (approximately 140 mOsm kg(-1)), phloem girdling had no effect on xylem hydraulic conductance. These results suggest a functional link between phloem and xylem hydraulic systems that is mediated by changes in the ionic content of the cell sap.     

Xylem cavitation and loss of hydraulic conductance in western hemlock following planting

Following planting, western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedlings experience water stress and declining xylem pressure potential (Psi(x)). Low Psi(x) can result in xylem cavitation and embolism formation, causing a decline in hydraulic conductance. This study focused on the relationship between Psi(x), xylem cavitation and transpiration (E) of newly planted seedlings. Leaf specific hydraulic conductance (k(AB)) declined from 0.56 to 0.09 mmol m(-2) s(-1) MPa(-1) over a 9-day period. Stomatal conductance (g(s)) declined from 143.5 to 39.15 mmol m(-2) s(-1) over the same period without an associated change in environmental conditions. A vulnerability profile indicated a 30% loss in hydraulic conductivity when seedlings experienced a Psi(x) between -2.5 and -3.0 MPa. A Psi(x) of -4.0 MPa led to a complete loss of conductivity. We conclude that following planting, western hemlock seedlings often experience Psi(x) values that are low enough to cause xylem cavitation and a decline in k(AB).     

Light response of hydraulic conductance in bur oak (Quercus macrocarpa) leaves

A four- to seven-fold enhancement of leaf hydraulic conductance by light has been reported in three temperate tree species. The enhancement occurs in the liquid-flow pathway between the petiole and the site of water evaporation. The enhancement occurs within 1 h, and dissipates in darkness over a period of 1 to 10 h depending on species. Here we report light-induced enhancement of leaf hydraulic conductance in a fourth species, bur oak (Quercus macrocarpa Michx.), the dependence of the effect on light flux and color, its absence in leaves of seedlings, and the impact on the response of leaf vein severance and several metabolic inhibitors. The light response of leaf hydraulic conductance approached saturation at a photosynthetic photon flux of 150 mumol m(-2) s(-1). Hydraulic enhancement was greater in response to blue and green light than to visible radiation of longer wavelengths, although at the same irradiance, the response to white light was greater than to light of any single color. Atrazine (a photosystem II inhibitor), fusicoccin (which stimulates plasma membrane-bound H(+)-ATPase) and HgCl(2) (an aquaporin blocker) reduced the light response of leaf lamina hydraulic conductance. When 2-mercaptoethanol was added following mercury treatment, the light response was totally suppressed. Our results are consistent with the notion that the effect of light on leaf lamina hydraulic conductance is controlled by factors acting outside the leaf veins, possibly through light-induced changes in membrane permeability of either mesophyll or bundle sheath cells, or both.     

Pit membrane chemistry influences the magnitude of ion-mediated enhancement of xylem hydraulic conductance in four Lauraceae species

The ion-mediated enhancement of xylem hydraulic conductivity in angiosperms is thought to be controlled by the pectin chemistry of intervessel pit membranes. However, there is little or no direct evidence on the ultrastructure and chemical nature of pit membranes in species that show an 'ionic effect'. The potential link between the magnitude of the ionic effect and pectin composition in intervessel pit membranes of four Lauraceae species (Laurus nobilis, Lindera megaphylla, Litsea sericea and Umbellularia californica) that show rather similar vessel and pit dimensions was studied using transmission electron microscopy (TEM). The TEM observations confirmed the presence of a pectic matrix associated with intervessel pit membranes, indicating that the relative abundance of acidic versus methylesterified pectins was closely related to the ionic effect. The two species examined with a high ionic effect ~20%, i.e. Laurus nobilis and Umbellularia californica) showed relatively high levels of acidic pectins, whereas methylesterified pectins were abundant in Lindera megaphylla and Litsea sericea, which showed a low ionic effect (~10%). Variation in the ionic effect is strongly associated with the chemical nature of pit membrane pectins in the species studied. Our findings support the current interpretation of the ionic effect due to dynamic swelling and shrinking behaviour of pit membrane pectins.     

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.     

Stomatal behavior of four woody species in relation to leaf-specific hydraulic conductance and threshold water potential

Midday stomatal closure is mediated by the availability of water in the soil, leaf and atmosphere, but the response to these environmental and internal variables is highly species specific. We tested the hypothesis that species differences in stomatal response to humidity and soil water availability can be explained by two parameters: leaf-specific hydraulic conductance (K(L)) and a threshold leaf water potential (Psi(threshold)). We used a combination of original and published data to estimate characteristic values of K(L) and Psi(threshold) for four common tree species that have distinctly different stomatal behaviors: black cottonwood (Populus trichocarpa Torr. & Gray.), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), red alder (Alnus rubra Bong.) and western hemlock (Tsuga heterophylla (Raf.) Sarg.). We used the values to parameterize a simple, nonelastic model that predicts stomatal conductance by linking hydraulic flux to transpirational flux and maintaining Psi(leaf) above Psi(threshold). The model successfully predicted fundamental features of stomatal behavior that have been reported in the literature for these species. We conclude that much of the variation among the species in stomatal response to soil and atmospheric water deficits can be explained by K(L) and Psi(threshold). The relationship between Psi(threshold) and xylem vulnerability to cavitation differed among these species.     

Fertilization effects on mean stomatal conductance are mediated through changes in the hydraulic attributes of mature Norway spruce trees

Stomatal conductance was quantified with sap flux sensors and whole-tree chambers in mature Norway spruce (Picea abies (L.) Karst.) trees after 3 years of exposure to elevated CO(2) concentration ([CO(2)]) in a 13-year nutrient optimization experiment. The long-term nutrient optimization treatment increased tree height by 3.7 m (67%) and basal diameter by 8 cm (68%); the short-term elevated [CO(2)] exposure had no effect on tree size or allometry. Nighttime transpiration was estimated as approximately 7% of daily transpiration in unchambered trees; accounting for the effect of nighttime flux on the processing of sap flux signals increased estimated daily water uptake by approximately 30%. Crown averaged stomatal conductance (g(s)) was described by a Jarvis-type model. The addition of a stomatal response time constant (tau) and total capacitance of stored water (C(tot)) improved the fit of the model. Model estimates for C(tot) scaled with sapwood volume of the bole in fertilized trees. Hydraulic support-defined as a lumped variable of leaf-specific hydraulic conductivity and water potential gradient (K(l)DeltaPsi) -was estimated from height, sapwood-to-leaf area ratio (A(s):A(l)) and changes in tracheid dimensions. Hydraulic support explained 55% of the variation in g(s) at reference conditions for trees across nutrient and [CO(2)] treatments. Removal of approximately 50% of A(l) from three trees yielded results suggesting that stomatal compensation (i.e., an increase in g(s)) after pruning scales inversely with K(l)DeltaPsi, indicating that the higher the potential hydraulic support after pruning, the less complete the stomatal compensation for the increase in A(s):A(l).     

Variations of root hydraulic conductance of Fraxinus mandshurica seedlings in different concentrations of NH4NO3 solution

Absorbing water from soil by roots in vascular plants is an important physiological function and plays an essential role on their water balance. The root hydraulic conductance (L _P) determined by radical water transport inside the root is a major influence on the shoot water status, plant growth, and development. However, a few studies have focused on the effect of different substances on L _P of roots, and the role of radical water transport was poorly understood. Based on the pressure-flux approach, this study used the roots of Fraxinus mandshurica seedlings with different treatments, i.e., distilled water, NH₄NO₃ solution, and HgCl₂ to determine the effect of various substances on L _P of roots. The objectives are: 1) to evaluate the difference in L _P occurred between distilled water and NH₄NO₃ solution with various concentrations; and 2) to examine the changes of L _P under distilled water and NH₄NO₃ solution with various concentrations after HgCl₂ treatment. The results showed that L _P of roots were 18.85×10⁻⁸ m/(s·MPa) in distilled water, 31.25–34.15×10⁻⁸ m/(s·MPa) in four NH₄NO₃ solutions (2, 4, 8 and 16 mmol/L), 14.69×10⁻⁸ m/(s·MPa) in distilled water after HgCl₂-treated, and 9.63–13.57×10⁻⁸ m/(s·MPa) in four NH₄NO₃ solutions after HgCl₂-treated, respectively. Aquaporins play an important role in regulating water uptake and transport in roots. NH₄ ⁺ and NO₃ ⁻ could stimulate activity of aquaporins, and L _P of roots in NH₄NO₃ solution was distinctly 77% higher than in distilled water. Nevertheless, Hg²⁺ can inhibit activity of aquaporins, and and L _P of roots decreased 22% in distilled water and 68% in NH₄NO₃ solution after treatment by HgCl₂ respectively. These evidences suggested that both Hg²⁺-sensitive aquaporins and ion channels existing in the protoplasm and vacuole membranes could regulate root water uptake, transport, and integral plant water balance. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(5): 706–712 [译自: 植物生态学报, 2005, 29(5): 706–712]     

Temperature, light and leaf hydraulic conductance of little-leaf linden (Tilia cordata) in a mixed forest canopy

Response of whole-leaf hydraulic conductance (G(L)) in little-leaf linden (Tilia cordata Mill.) to temperature and photosynthetic photon flux (Q(P)) was estimated by the evaporative flux method under natural conditions in a mixed forest canopy. Mean midday G(L) in the lower- and upper-crown foliage was 1.14 and 3.06 mmol m(-2) s(-1) MPa(-1), respectively. Over the study period, leaf temperature (T(L)) explained about 67% of the variation in G(L), and Q(P) explained about 10%. Leaf water potential and crown position also affected G(L) significantly. About a third of the temperature effect was attributable to changes in the viscosity of water, and two thirds to changes in protoplast permeability (i.e., symplastic conductance). Leaf hydraulic conductance was highly sensitive to changes in Q(P) when Q(P) was less than 200 micromol m(-2) s(-1), and G(L) sensitivity decreased with increasing irradiance. Sensitivity of G(L) to variation in T(L) increased consistently with increasing temperature in the range of 16 to 29 degrees C. There were positive interactions between temperature and light in their effects on G(L): the light response was more pronounced at higher leaf temperatures. Because of frequent rains during the study period, the trees experienced no soil water deficit, and, within the range experienced, soil water potential had no effect on G(L). Leaf hydraulic conductance exhibited a seasonal pattern that could be explained primarily by temporal variability in mean air temperature and irradiance, in addition to which an age-related trend (P<0.001) of increasing G(L) from the end of June to the beginning of August was observed.     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

Recovery of diurnal depression of leaf hydraulic conductance in a subtropical woody bamboo species: embolism refilling by nocturnal root pressure

Despite considerable investigations of diurnal water use characteristics in different plant functional groups, the research on daily water use strategies of woody bamboo grasses remains lacking. We studied the daily water use and gas exchange of Sinarundinaria nitida (Mitford) Nakai, an abundant subtropical bamboo species in Southwest China. We found that the stem relative water content (RWC) and stem hydraulic conductivity (K(s)) of this bamboo species did not decrease significantly during the day, whereas the leaf RWC and leaf hydraulic conductance (K(leaf)) showed a distinct decrease at midday, compared with the predawn values. Diurnal loss of K(leaf) was coupled with a midday decline in stomatal conductance (g(s)) and CO(2) assimilation. The positive root pressures in the different habitats were of sufficient magnitude to refill the embolisms in leaves. We concluded that (i) the studied bamboo species does not use stem water storage for daily transpiration; (ii) diurnal down-regulation in K(leaf) and gs has the function to slow down potential water loss in stems and protect the stem hydraulic pathway from cavitation; (iii) since K(leaf) did not recover during late afternoon, refilling of embolism in bamboo leaves probably fully depends on nocturnal root pressure. The embolism refilling mechanism by root pressure could be helpful for the growth and persistence of this woody monocot species.     

Effect of galls induced by Endocronartium harknessii on stem hydraulic conductivity and growth of lodgepole pine

To test the hypothesis that galls resulting from infection by western gall rust [Endocronartium harknessii (J. P. Moore) Y. Hiratsuka] on lodgepole pine [Pinus contorta Dougl. ex Loud.] create hydraulic constrictions that reduce tree growth, the hydraulic conductivity (K_ψ; m² s^?1 Pa^?1), leaf area (A_L) and sapwood area (A_S) were measured on 6‐ to 15‐year‐old trees having galls encircling 0–100% of the stem. Increases in gall encirclement were accompanied by decreases in K_ψ, foliar nitrogen concentration, A_L and A_S, although the A_L:A_S ratio remained constant. Dye flow patterns through galled stem sections indicated that water moved primarily through the non‐galled portions of the stem. In a second field study, xylem and phloem wounds were applied to 13‐year‐old galled and non‐galled trees on either the galled or non‐galled side of the tree. Although both galls and xylem wounds reduced K_ψ, radial and height growth were unaffected. In aggregate the two experiments showed that although trees are quite tolerant to small galls and short term reductions in K_ψ, the disruption of water movement by larger galls may be sufficient to initiate a chain of events leading to tree death, especially in very dense stands with high levels of intraspecific competition.     

Predicting the decline in daily maximum transpiration rate of two pine stands during drought based on constant minimum leaf water potential and plant hydraulic conductance

The effect of drought on forest water use is often estimated with models, but comprehensive models require many parameters, and simple models may not be sufficiently flexible. Many tree species, Pinus species in particular, have been shown to maintain a constant minimum leaf water potential above the critical threshold for xylem embolism during drought. In such cases, prediction of the relative decline in daily maximum transpiration rate with decreasing soil water content is relatively straightforward. We constructed a soil-plant water flow model assuming constant plant conductance and daily minimum leaf water potential, but variable conductance from soil to root. We tested this model against independent data from two sites: automatic shoot chamber data and sap flow measurements from a boreal Scots pine (Pinus sylvestris L.) stand; and sap flow measurements from a maritime pine (Pinus pinaster Ait.) stand. To focus on soil limitations to water uptake, we expressed daily maximum transpiration rate relative to the rate that would be obtained in wet soil with similar environmental variables. The comparison was successful, although the maritime pine stand showed carry-over effects of the drought that we could not explain. For the boreal Scots pine stand, daily maximum transpiration was best predicted by water content of soil deeper than 5 cm. A sensitivity analysis revealed that model predictions were relatively insensitive to the minimum leaf water potential, which can be accounted for by the importance of soil resistance of drying soil. We conclude that a model with constant plant conductance and minimum leaf water potential can accurately predict the decline in daily maximum transpiration rate during drought for these two pine stands, and that including further detail about plant compartments would add little predictive power, except in predicting recovery from severe drought.     

Effects of summer drought and winter freezing on stem hydraulic conductivity of Rhododendron species from contrasting climates

We studied the limits to maximum water transport in three diffuse-porous evergreen shrubs exposed to frequent winter freeze-thaw events (Rhododendron maximum L. and R. catawbiense Michaux from the Appalachian Mountains) and to a severe summer drought (R. macrophyllum G. Don. from the Oregon Cascades). Percent loss of hydraulic conductivity (PLC), vulnerability curves to xylem embolism and freezing point temperatures of stems were measured over 2 years. Controlled freeze-thaw experiments were also conducted to determine the effect of thaw rate on PLC. During both years, native PLC was significantly higher in winter than in summer for R. macrophyllum. Seasonal changes in PLC were variable in both R. catawbiense and R. maximum. Only R. maximum plants growing in gaps or clearings showed higher PLC than understory plants. A rapid (2-4 day) natural recovery of high native PLC during the winter was observed in both R. maximum and R. macrophyllum. Compared with the bench-dehydration method, vulnerability curves based on the air-injection method consistently had less negative slopes and greater variation. Fifty percent PLC (PLC(50)) obtained from vulnerability curves based on the dehydration method occurred at -1.75, -2.42 and -2.96 MPa for R. catawbiense, R. maximum and R. macrophyllum, respectively. Among the study species, R. macrophyllum, which commonly experiences a summer drought, had the most negative water potential at PLC(50). In all species, stem freezing point temperatures were not consistently lower in winter than in summer. A single fast freeze-thaw event significantly increased PLC, and R. catawbiense had the highest PLC in response to freezing treatments. Recovery to control PLC values occurred if a low positive hydraulic pressure was maintained during thawing. Rhododendron macrophyllum plants, which commonly experience few freeze-thaw events, had large stem diameters, whereas plants of R. catawbiense, which had small stem diameters, suffered high embolism in response to a single freeze-thaw event. Both drought-induced and winter-induced embolism caused a significant reduction in hydraulic conductivity in all species during periods when drought or freeze-thaw events occurred in their native habitats. However, rapid recovery of PLC following freezing or drought maintained the species above their relatively low margins of safety for complete xylem dysfunction.     

Effects of ring-porous and diffuse-porous stem wood anatomy on the hydraulic parameters used in a water flow and storage model

Calibration of a recently developed water flow and storage model based on experimental data for a young diffuse-porous beech tree (Fagus sylvatica L.) and a young ring-porous oak tree (Quercus robur L.) revealed that differences in stem wood anatomy between species strongly affect the calibrated values of the hydraulic model parameters. The hydraulic capacitance (C) of the stem storage tissue was higher in oak than in beech (939.8 versus 212.3 mg MPa(-1)). Model simulation of the elastic modulus (epsilon) revealed that this difference was linked to the higher elasticity of the stem storage tissue of oak compared with beech. Furthermore, the hydraulic resistance (R (x)) of beech was about twice that of oak (0.1829 versus 0.1072 MPa s mg(-1)). To determine the physiological meaning of the R (x) parameter identified by model calibration, we analyzed the stem wood anatomy of the beech and oak trees. Calculation of stem specific hydraulic conductivity (k (s)) of beech and oak with the Hagen-Poiseuille equation confirmed the differences in R (x) predicted by the model. The contributions of different vessel diameter classes to the total hydraulic conductivity of the xylem were calculated. As expected, the few big vessels contributed much more to total conductivity than the many small vessels. Compared with beech, the larger vessels of oak resulted in a higher k (s) (10.66 versus 4.90 kg m(-1) s(-1) MPa(-1)). The calculated ratio of k (s) of oak to beech was 2, confirming the R (x) ratio obtained by model calibration. Thus, validation of the R (x) parameter of the model led to identification of its physiological meaning.     

Experimental evidence for the relative sensitivity of deciduous forest plants to high soil acidity

Thirteen species of deciduous forest vascular plants were established from seeds: Brachypodium sylvaticum, Carex sylvatica, Circaea lutetiana, Deschampsia flexuosa, Galium odoratum, Lamium galeobdolon, Lathyrus vernus, Luzula pilosa, Milium effusum, Poa nemoralis, Sanicula europaea, Stellaria holostea and Stellaria nemorum. An acid beech forest soil was variously treated with carbonates (CaCO₃ + MgCO₃ or SrCO₃) to obtain a soil solution pH of 3.3–5.2. The experimental results were compared with the field distribution of the same species in 230 south Swedish deciduous forests with regard to soil pH.

In general, the species reacted to increasing soil pH in accordance with their field distribution. Addition of SrCO₃ was almost equally effective as CaCO₃ + MgCO₃ (both at pH 4.2 in the soil solution) in promoting establishment and growth, whereas most species failed in the untreated soil (pH 3.3). There were no indications that the Ca or Mg concentrations of the soil solution were below minimum or even suboptimal for plant growth in the untreated soil. Within the pH range (pH 3.5–4.3) which proved critical to establishment and growth in most of the species, the reactive Al³⁺ concentration of the soil solution did not vary significantly. It is suggested that the high acidity of the rhizosphere solution, possibly aggravated by Al³⁺, obstructed the uptake or retention of mineral nutrients by the roots in the untreated acid beech forest soil.     

Rate of stomatal opening, shoot hydraulic conductance and photosynthetic characteristics in relation to leaf abscisic acid concentration in six temperate deciduous trees

Correlations between leaf abscisic acid concentration ([ABA]), stomatal conductance (gs), rate of stomatal opening in response to an increase in leaf water potential (si), shoot hydraulic conductance (L) and photosynthetic characteristics were examined in saplings of six temperate deciduous tree species: Acer platanoides L., Padus avium Mill., Populus tremula L., Quercus robur L., Salix caprea L. and Tilia cordata Mill. Species-specific values of foliar [ABA] were negatively related to the mean values of gs, si, L and light- and CO2- saturated net photosynthesis (P(max)), thus providing strong correlative evidence of a scaling of foliar gas exchange and hydraulic characteristics with leaf endogenous [ABA]. In addition, we suggest that mean gs, si, L and Pmax for mature leaves may partly be determined by the species-specific [ABA] during leaf growth. The most drought-intolerant species had the lowest [ABA] and the highest gs, suggesting that interspecific differences in [ABA] may be linked to differences in species-specific water-use efficiency. Application of high concentrations of exogenous ABA led to large decreases in gs, si and P(max), further underscoring the direct role of ABA in regulating stomatal opening and photosynthetic rate. Exogenous ABA also decreased L, but the decreases were considerably smaller than the decreases in gs, si and Pmax. Thus, exogenous ABA predominantly affected the stomata directly, but modification of L by ABA may also be an important mechanism of ABA action. We conclude that interspecific variability in endogenous [ABA] during foliage growth and in mature leaves provides an important factor explaining observed differences in L, gs, si and Pmax among temperate deciduous tree species.     

Effects of drought stress and high density stem inoculations with Leptographium wingfieldii on hydraulic properties of young Scots pine trees

We examined drought-induced changes in susceptibility of potted Scots pine (Pinus sylvestris L.) trees to a bark-beetle associated fungus (Leptographium wingfieldii Morelet, from the bark beetle Tomicus piniperda L.). Five-year-old field-grown trees were transplanted to 50-l pots and grown for 1 year before the treatments were applied. Trees in the drought treatment were subjected to several successive, 3-week-long drought cycles, with predawn water potential dropping below -2 MPa at peak drought intensity. The experimental drought cycles were more severe than the natural drought episodes usually recorded in Scots pine stands. Trees were then mass-inoculated with L. wingfieldii at a density close to the critical threshold density of inoculations (400 m(-2)) above which tree resistance is overcome. Inoculation of well-watered trees resulted in induced reaction zones around the inoculation points and very limited damage (resinosis) in the sapwood. Drought alone had no long-lasting consequences on tree water relations, except for a decrease in hydraulic conductance in the youngest segments of the main stem. However, the combination of mass-inoculation and drought stress after inoculation resulted in a dramatic loss of stem hydraulic conductivity that was paralleled by conspicuous damage to the sapwood (resinosis, drying and blue staining). There was a close correlation between amount of visible sapwood damage and loss of hydraulic conductivity. The intensity of induced reactions in the phloem was unaffected by drought stress. We conclude that tree defence against L. wingfieldii is decreased during severe drought stress, resulting in changes in the spread and action of the fungus in the sapwood but not in the phloem.