Hydraulic efficiency and safety of vascular and non-vascular components in Pinus pinaster leaves

Leaves, the distal section of the soil-plant-atmosphere continuum, exhibit the lowest water potentials in a plant. In contrast to angiosperm leaves, knowledge of the hydraulic architecture of conifer needles is scant. We investigated the hydraulic efficiency and safety of Pinus pinaster needles, comparing different techniques. The xylem hydraulic conductivity (k(s)) and embolism vulnerability (P(50)) of both needle and stem were measured using the cavitron technique. The conductance and vulnerability of whole needles were measured via rehydration kinetics, and Cryo-SEM and 3D X-ray microtomographic observations were used as reference tools to validate physical measurements. The needle xylem of P. pinaster had lower hydraulic efficiency (k(s)?=?2.0?×?10(-4) m(2) MPa(-1) s(-1)) and safety (P(50)?=?-?1.5 MPa) than stem xylem (k(s)?=?7.7?×?10(-4) m(2) MPa(-1) s(-1); P(50)?=?-?3.6 to?-?3.2 MPa). P(50) of whole needles (both extra-vascular and vascular pathways) was?-?0.5 MPa, suggesting that non-vascular tissues were more vulnerable than the xylem. During dehydration to?-?3.5 MPa, collapse and embolism in xylem tracheids, and gap formation in surrounding tissues were observed. However, a discrepancy in hydraulic and acoustic results appeared compared with visualizations, arguing for greater caution with these techniques when applied to needles. Our results indicate that the most distal parts of the water transport pathway are limiting for hydraulics of P. pinaster. Needle tissues exhibit a low hydraulic efficiency and low hydraulic safety, but may also act to buffer short-term water deficits, thus preventing xylem embolism.     

Water relations and drought-induced embolism in olive (Olea europaea) varieties 'Meski' and 'Chemlali' during severe drought

We examined the effects of drought on the water relations, osmotic adjustment and xylem vulnerability to embolism of olive (Olea europaea L.) varieties, 'Meski' and 'Chemlali'. Two-year-old self-rooted cuttings growing in sand-filled pots in a greenhouse were subjected to water stress by withholding water for 60 days. Water relations and gas exchange measurements showed that 'Chemlali' was more drought resistant than 'Meski' and had a greater capacity for osmotic adjustment through solute accumulation. However, when water stress was acute, the effect of osmoregulation on leaf cell turgor was largely counteracted by xylem cavitation. Cavitation vulnerability curves showed that both varieties were highly resistant to embolism formation. The xylem water potential inducing 50% loss of stem conductivity approached -7 MPa in 'Meski' and only slightly less in 'Chemlali'. Although the difference between varieties in susceptibility to xylem embolism was small, it appears to account in large part for the difference between them in the ability to tolerate severe drought.     

Patterns of hydraulic architecture and water relations of two tropical canopy trees with contrasting leaf phenologies: Ochroma pyramidale and Pseudobombax septenatum

Many authors have attempted to explain the adaptive response of tropical plants to drought based on studies of water relations at the leaf level. Little attention has been given to the role of the xylem system in the control of plant water requirements. To evaluate this role, we studied the hydraulic architecture and water relations parameters of two tropical canopy trees with contrasting leaf phenologies: deciduous Pseudobombax septenatum (Jacq.) Dug and evergreen Ochroma pyramidale (Cav. ex lamb) Urban, both in the family Bombacaceae. The hydraulic architecture parameters studied include hydraulic conductivity, specific conductivity, leaf specific conductivity, and Huber value. Water relations parameters include leaf water potential, stem and leaf water storage capacitance, transpiration, stomatal conductance, and vulnerability of stems to cavitation and loss of hydraulic conductivity by embolisms. Compared to temperate trees, both species showed a pattern of highly vulnerable stems (50% loss of conductivity due to embolism at water potentials less than 1 MPa) with high leaf specific conductivities. The vulnerability of xylem to water-stress-induced embolism was remarkably similar for the two species but the leaf specific conductivity of petioles and leaf-bearing stems of the evergreen species, Ochroma (e.g., 9.08 and 11.4 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively), were 3.4 and 2.3 times higher, respectively, than those of the deciduous species, Pseudobombax (e.g., 2.64 and 5.15 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively). A runaway embolism model was used to test the ability of Ochroma and Pseudobombax stems to maintain elevated transpiration rates during the higher evaporative demand of the dry season. The percent loss of leaf area predicted by the runaway embolism model for stems of Pseudobombax ranged from 5 to 30%, not enough to explain the deciduous phenology of this tree species without analysis of root resistance or leaf and petiole vulnerability to embolism.     

Formation and seasonal occurrence of xylem embolism in Alnus cordata

We investigated the vulnerability of xylem to embolism and the seasonal occurrence of xylem embolism in Italian alder (Alnus cordata Loisel.) by acoustic and hydraulic methods. Wood anatomy was also studied. More than eighty percent of the vessels were less than 50 mm long and no vessels were longer than 120 mm. Mean vessel diameter was 48 micro m. Ultrasound acoustic emissions from root and branch segments dehydrating in air followed a similar pattern: in both tissues, emission peaks were recorded when the relative water content of the xylem was around 0.2. In branches dehydrating in air, xylem embolism increased linearly as water potential decreased. In trees in the field, more than 80 percent of hydraulic conductivity was lost in the tree crowns during winter. Recovery from winter embolism occurred mostly before bud burst. In summer, xylem embolism was low (< 30%) and acoustic emissions from roots, stem and branches of trees in the field were also low.     

Drought effects on hydraulic conductivity and xylem vulnerability to embolism in diverse species and provenances of Mediterranean cedars

We studied hydraulic traits of young plants of the Mediterranean cedar species Cedrus atlantica (Endl.) G. Manetti ex Carrière (Luberon, France), C. brevifolia (Hook. f.) Henry (Cyprus), C. libani A. Rich (Hadeth El Jebbe, Lebanon) and C. libani (Armut Alani, Turkey). With an optimum water supply, no major differences were observed among species or provenances in either stem hydraulic conductivity (Ks) or leaf specific conductivity (Kl) measured on the main shoot. A moderate soil drought applied for 10 weeks induced marked acclimation through a reduction in Ks, particularly in the Lebanese provenance of C. libani, and a decrease in tracheid lumen size in all species. Cedrus atlantica, which had the smallest tracheids, was the species most vulnerable to embolism: a 50% loss in hydraulic conductivity (PsiPLC50) occurred at a water potential of -4.4 MPa in the well-watered treatment, and at -6.0 MPa in the moderate drought treatment. In the other species, PsiPLC50 was unaffected by moderate soil drought, and only declined sharply at water potentials between -6.4 and -7.5 MPa in both irrigation treatments. During severe drought, Ks of twigs and stomatal conductance (g(s)) were measured simultaneously as leaf water potential declined. For all species, lower vulnerability to embolism based on loss of Ks was recorded on current-year twigs. The threshold for stomatal closure (10% of maximum g(s)) was reached at a predawn water potential (Psi(pd)) of -2.5 MPa in C. atlantica (Luberon) and at -3.1 MPa in C. libani (Lebanon), whereas the other provenance and species had intermediate Psi(pd) values. Cedrus brevifolia, with a Psi(pd) (-3.0 MPa) close to that of C. libani (Lebanon), had the highest stomatal conductance of the study species. The importance of a margin of safety between water potential causing stomatal closure and that causing xylem embolism induction is discussed.     

Interspecific variation in xylem vulnerability to cavitation among tropical tree and shrub species

In tropical moist forests, seasonal drought limits plant survival, productivity and diversity. Drought-tolerance mechanisms of tropical species should reflect the maximum seasonal water deficits experienced in a particular habitat. We investigated stem xylem vulnerability to cavitation in nine tropical species with different life histories and habitat associations. Stem xylem vulnerability was scored as the xylem water potential causing 50 and 75% loss of hydraulic conductivity (P50 and P75, respectively). Four shade-tolerant shrubs ranged from moderately resistant (P50=-1.9 MPa for Ouratea lucens Kunth. Engl.) to highly resistant to cavitation (P50=-4.1 MPa for Psychotria horizontalis Sw.), with shallow-rooted species being the most resistant. Among the tree species, those characteristic of waterlogged soils, Carapa guianensis Aubl., Prioria copaifera Griseb. and Ficus citrifolia Mill., were the most vulnerable to cavitation (P50=-0.8 to -1.6 MPa). The wet-season, deciduous tree, Cordia alliodora (Ruiz and Pav.) Oken., had resistant xylem (P50=-3.2 MPa), whereas the dry-season, deciduous tree, Bursera simaruba (L.) Sarg. was among the most vulnerable to cavitation (P50=-0.8 MPa) of the species studied. For eight out of the nine study species, previously reported minimum seasonal leaf water potentials measured in the field during periods of drought correlated with our P50 and P75 values. Rooting depth, deciduousness, soil type and growth habit might also contribute to desiccation tolerance. Our results support the functional dependence of drought tolerance on xylem resistance to cavitation.     

Loss of hydraulic conductivity due to water stress in intact juveniles of Quercus rubra and Populus deltoides

Despite many studies of the percent loss of hydraulic conductivity in excised branches, there is doubt as to whether cutting stems in air introduces unnatural embolism into the xylem at the cut surface. To address this question, hydraulic conductivity was measured in seedlings of northern red oak (Quercus rubra L.) and rooted scions of eastern cottonwood (Populus deltoides Bartr. ex Marsh.) that had been droughted in pots. Results indicate that in situ dehydration produced a very similar vulnerability curve (% loss of conductivity versus water potential) to those previously obtained by bench-top dehydration of excised branches of eastern cottonwood and red oak. In eastern cottonwood cuttings, conductivity loss increased sharply below water potentials of -1.0 MPa, with 100% loss of conductivity occurring by -2.0 MPa, whereas conductivity loss in red oak seedlings was more gradual, i.e., increasing below -1.5 MPa and sustaining 100% loss of conductivity by about -4.0 MPa.     

Whole-plant hydraulic resistance and vulnerability segmentation in Acer saccharinum

Hydraulic properties were studied in Acer saccharinum L., a riparian species that also grows well on a dry soil when transplanted. Hydraulic resistances were measured by two independent techniques: a new high-pressure flowmeter (HPFM) method and a conventional evaporative flux (EF) method. Vulnerability to cavitation was also investigated on petioles, stems and roots using a hydraulic conductivity technique. Vulnerability segmentation was found, i.e., roots, stems and petioles had different vulnerabilities to xylem dysfunction. Petioles were most vulnerable with 50% loss of hydraulic conductivity at -0.5 MPa, roots were least vulnerable (50% loss at -2.2 MPa) and stems were intermediate in vulnerability. The HPFM and the EF methods gave comparable results, except that the EF method gave a significantly higher value for resistance across petioles plus leaves. Native embolism was high enough to explain the discrepancy in resistance across petioles plus leaves between the HPFM and the EF methods, indicating that the HPFM estimates the minimum (potential) hydraulic resistance of plants. Whole-plant hydraulic resistance of A. saccharinum was low compared to resistances of other temperate species. The hydraulic characteristics of A. saccharinum were consistent with adaptation to its typical environment: low whole-plant resistance assures high transpiration rates in the presence of sufficient water, and vulnerability segmentation provides the ability to survive during droughts through shedding of expendable organs.     

Cavitation in dehydrating xylem of Picea abies: energy properties of ultrasonic emissions reflect tracheid dimensions

Ultrasonic emission (UE) testing is used to analyse the vulnerability of xylem to embolism, but the number of UEs often does not sufficiently reflect effects on hydraulic conductivity. We monitored the absolute energy of UE signals in dehydrating xylem samples hypothesizing that (i) conduit diameter is correlated with UE energy and (ii) monitoring of UE energy may enhance the utility of this technique for analysis of xylem vulnerability. Split xylem samples were prepared from trunk wood of Picea abies, and four categories of samples, derived from mature (I: earlywood, II: 30-50% latewood, III: >50% latewood) or juvenile wood (IV: earlywood) were used. Ultrasonic emissions during dehydration were registered and anatomical parameters (tracheid lumen area, number per area) were analysed from cross-sections. Attenuation of UE energy was measured on a dehydrating wood beam by repeated lead breaks. Vulnerability to drought-induced embolism was analysed on dehydrating branches by hydraulic, UE number or UE energy measurements. In split samples, the cumulative number of UEs increased linearly with the number of tracheids per cross-section, and UE energy was positively correlated with the mean lumen area. Ultrasonic emission energies of earlywood samples (I and IV), which showed normally distributed tracheid lumen areas, increased during dehydration, whereas samples with latewood (II and III) exhibited a right-skewed distribution of lumina and UE energies. Ultrasonic emission energy was hardly influenced by moisture content until ～40% moisture loss, and decreased exponentially thereafter. Dehydrating branches showed a 50% loss of conductivity at -3.6 MPa in hydraulic measurements and at -3.9 and -3.5 MPa in UE analysis based on cumulative number or energy of signals, respectively. Ultrasonic emission energy emitted by cavitating conduits is determined by the xylem water potential and by the size of element. Energy patterns during dehydration are thus influenced by the vulnerability to cavitation, conduit size distribution as well as attenuation properties. Measurements of UE energy may be used as an alternative to the number of UEs in vulnerability analysis.     

Hydraulic conductivity and embolism in the mangrove tree Laguncularia racemosa

We measured xylem pressure potentials, soil osmotic potentials, hydraulic conductivity and percent loss of conductivity (PLC) due to embolism, and made microscopic observations of perfused dye in the white mangrove tree, Laguncularia racemosa (L.) Gaertn. f., (1) to determine its vulnerability to air embolism compared with published results for the highly salt-tolerant red mangrove tree, Rhizophora mangle L., and (2) to identify possible relationships between air embolism, permanent blockage of vessels and stem diameter. Laguncularia racemosa was more vulnerable to embolism than reported for R. mangle, with 50 PLC at -3.4 MPa. Narrow stems (5-mm diameter) had higher PLC than larger stems (8.4- or 14-mm diameter) of the same plants. Basic fuchsin dye indicated that up to 89% of the vessels, especially in the narrow stems, had permanent blockage that could not be reversed by high pressure perfusion. Air embolism could lead to permanent vessel blockage and eventual stem mortality. Such vulnerability to embolism may restrict the growth of L. racemosa and limit its distribution to less salty areas of mangrove communities.     

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.     

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).     

An inconvenient truth about xylem resistance to embolism in the model species for refilling <Emphasis Type="Italic">Laurus nobilis</Emphasis> L.

Key message

Direct, non-invasive X-ray microtomography and optical technique observations applied in stems and leaves of intact seedlings revealed that laurel is highly resistant to drought-induced xylem embolism. Contrary to what has been brought forward, daily cycles of embolism formation and refilling are unlikely to occur in this species and to explain how it copes with drought.

Context

There has been considerable controversy regarding xylem embolism resistance for long-vesselled angiosperm species and particularly for the model species for refilling (Laurus nobilis L.).

Aims

The purpose of this study was to resolve the hydraulic properties of this species by documenting vulnerability curves of different organs in intact plants.

Methods

Here, we applied a direct, non-invasive method to visualize xylem embolism in stems and leaves of intact laurel seedlings up to 2-m tall using X-ray microtomography (microCT) observations and the optical vulnerability technique. These approaches were coupled with complementary centrifugation measurements performed on 1-m long branches sampled from adult trees and compared with additional microCT analyses carried out on 80-cm cut branches.

Results

Direct observations of embolism spread during desiccation of intact laurels revealed that 50% loss of xylem conductivity (Ψ₅₀) was reached at ??7.9?±?0.5 and ??8.4?±?0.3 MPa in stems and leaves, respectively, while the minimum xylem water potentials measured in the field were ??4.2 MPa during a moderate drought season. Those findings reveal that embolism formation is not routine in Laurus nobilis contrary to what has been previously reported. These Ψ₅₀ values were close to those based on the flow-centrifuge technique (??9.2?±?0.2 MPa), but at odds with microCT observations of cut branches (??4.0?±?0.5 MPa).

Conclusion

In summary, independent methods converge toward the same conclusion that laurel is highly resistant to xylem embolism regardless its development stage. Under typical growth conditions without extreme drought events, this species maintains positive hydraulic safety margin, while daily cycles of embolism formation and refilling are unlikely to occur in this species.

     

Shoot and root vulnerability to xylem cavitation in four populations of Douglas-fir seedlings

The objectives of this study were to assess the range of genotypic variation in the vulnerability of the shoot and root xylem of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings to water-stress-induced cavitation, and to assess the trade-off between vulnerability to cavitation and conductivity per unit of stem cross-sectional area (k(s)), both within a species and within an individual tree. Douglas-fir occupies a broad range of environments and exhibits considerable genetic variation for growth, morphology, and drought hardiness. We chose two populations from each of two varieties (the coastal var. menziesii and the interior var. glauca) to represent environmental extremes of the species. Vulnerability curves were constructed for shoots and roots by plotting the percentage loss in conductivity versus water potential. Vulnerability in shoot and root xylem varied genetically with source climate. Stem xylem differed in vulnerability to cavitation between populations; the most mesic population, coastal wet (CW), was the most susceptible of the four populations. In the roots, the most vulnerable population was again CW; the interior wet (IW) population was moderately susceptible compared with the two dry populations, coastal dry (CD) and interior dry (ID). Root xylem was more susceptible to cavitation than stem xylem and had significantly greater k(s). The trade-off between vulnerability to cavitation and k(s), however, was not evident across populations. The most vulnerable population (CW) had a shoot k(s) of 0.534 +/- 0.067 &mgr;mol m(-2) s(-1) MPa(-1), compared with 0.734 +/- 0.067 &mgr;mol m(-2) s(-1) MPa(-1) for the less vulnerable CD stems. In the roots, IW was more vulnerable than ID, but had the same k(s).     

Xylem conductivity and vulnerability to cavitation of ponderosa pine growing in contrasting climates

We examined the effects of increased transpiration demand on xylem hydraulic conductivity and vulnerability to cavitation of mature ponderosa pine (Pinus ponderosa Laws.) by comparing trees growing in contrasting climates. Previous studies determined that trees growing in warm and dry sites (desert) had half the leaf/sapwood area ratio (A(L)/A(S)) and more than twice the transpiration rate of trees growing in cool and moist sites (montane). We predicted that high transpiration rates would be associated with increased specific hydraulic conductivity (K(S)) and increased resistance to xylem cavitation. Desert trees had 19% higher K(S) than montane trees, primarily because of larger tracheid lumen diameters. Predawn water potential and water potential differences between the soil and the shoot were similar for desert and montane trees, suggesting that differences in tracheid anatomy, and therefore K(S), were caused primarily by temperature and evaporative demand, rather than soil drought. Vulnerability to xylem cavitation did not differ between desert and montane populations. A 50% loss in hydraulic conductivity occurred at water potentials between -2.61 and -2.65 MPa, and vulnerability to xylem cavitation did not vary with stem size. Minimum xylem tensions of desert and montane trees did not drop below -2.05 MPa. Foliage turgor loss point did not differ between climate groups and corresponded to mean minimum xylem tensions in the field. In addition to low A(L)/A(S), high K(S) in desert trees may provide a way to increase tree hydraulic conductivity in response to high evaporative demand and prevent xylem tensions from reaching values that cause catastrophic cavitation. In ponderosa pine, the flexible responses of A(L)/A(S) and K(S) to climate may preclude the existence of significant intraspecific variation in the vulnerability of xylem to cavitation.     

Hydraulic architecture and water relations of a flood-tolerant tropical tree, Annona glabra

Hydraulic architecture parameters, water relation parameters and wood anatomy were studied in roots and shoots of the flood-tolerant tree Annona glabra L. on Barro Colorado Island, Panama. Hydraulic conductivity, leaf specific conductivity, and Huber value were similar to the corresponding values for tree species living in non-flooded habitats. The vulnerability of stems to loss of hydraulic conductivity resulting from embolism was low (50% loss of conductivity at -3.3 MPa). The lowest leaf water potential measured in the field was about -1.0 MPa, indicating that A. glabra has a large margin of safety from embolism, which may provide protection against rare drought events, or may be an adaptation to brackish mangrove habitats. Low absolute conductivity of roots was compensated for by an increase in the number of roots. More than two-thirds of whole-plant resistance to water flow was located in the roots.     

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.     

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.     

Development and recovery from winter embolism in silver birch: seasonal patterns and relationships with the phenological cycle in oceanic Scotland

Silver birch (Betula pendula Roth) is increasingly used in the United Kingdom for reforestation. However, recent evidence indicates that, under some circumstances, planted birch can suffer serious and repeated mortality of the apical leaders and branches, with consequent loss of apical dominance and the formation of a contorted stem. Plants from 37 seed sources of silver birch from Scotland and northern England planted at two sites were compared for several characteristics related to hydraulic architecture, vulnerability to freeze-thaw cycle induced embolism and spring recovery from winter embolism during the period 2000-2002. Phenological rhythms were also monitored in late winter-early spring to document relationships between phenology and water relations parameters. Significant differences were found across seed sources in stage of bud flushing for four dates in spring. Early flushing seed sources differed by about 1 to 2 weeks from late-flushing seed sources across the two sites. Wintertime xylem embolism in stems reached a peak of about 50 to 70% loss of xylem hydraulic conductivity, depending on the size and position of the sample shoots in the canopy. Small apical shoots were significantly more embolized than large basal shoots. Development of winter embolism was coupled to the occurrence of frost events. As percent loss of hydraulic conductivity increased during the winter, wood relative water content declined. Embolism reversal occurred rapidly in spring at the time of development of positive root pressure. No significant differences in the degree of winter embolism in 2001 were found among the three seed sources examined. The investigation was expanded in the winter-spring of 2002 to include 10 seed sources across both sites. Significant differences were found in degree of winter embolism across sites, dates and seed sources. For each date, there was a significant relationship between flushing scores and wood relative water contents across the two sites and all seed sources, suggesting that differences in time of flushing across sites and seed sources were likely caused by differences in the time of occurrence of root pressure, a necessary precondition to flushing.     

A Discussion on Drought Tolerance Through Hydraulic Architecture of Woody Plants

In this paper, ten woody plants grew on the campus of the Beijing Forestry University were selected for measuring hydraulic architecture parameters and water potential of one-year-old twigs. The results show that day and night change of water potential and hydraulic architecture parameters appear to be obviously convex trend. The relationship models between water potential and hydraulic conductivity, special conductivity and leaf special conductivity were respectively established, which were simulated by equations Y=axb and Y=ax2 bx c, through which the parameters physiological meanings were discussed. The results show also show that embolism vulnerability of ten tree species was Robinia pseudoacacia> Gleditsia sinensis> Acer truncatum> Prunus davidiana> Cornus alba> Platycladus orientalis> Cedrus deodara> Pinus bungeana> Pinus tabulaeformis> Jasminum nudiflorum, and that the water transport efficiency in xylem and embolism vulnerability of the species are ring-porous species> diffuse-porous species>non