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.     

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).     

Vulnerability to drought-induced cavitation of riparian cottonwoods in Alberta: a possible factor in the decline of the ecosystem?

Vulnerability of xylem to loss of hydraulic conductivity caused by drought-induced cavitation was determined for three riparian cottonwood species in Lethbridge, Alberta: Populus deltoides Bartr., P. balsamifera L., and P. angustifolia James. These species suffered 50% loss of hydraulic conductivity in one-year-old stem segments when xylem pressure potential fell to -0.7 MPa for P. deltoides and -1.7 MPa for P. balsamifera and P. angustifolia, making them the three most vulnerable tree species reported so far in North America. The possible contribution of drought-induced xylem dysfunction to the decline of riparian ecosystems in dammed rivers is discussed.     

Regulation of water loss in populations of Populus trichocarpa: the role of stomatal control in preventing xylem cavitation

Variations in resistance to drought-induced xylem cavitation, xylem air-entry points, stomatal behavior, and hydraulic conductivity were measured in four populations of Populus trichocarpa Torr. & A. Gray collected along an east-west humidity and temperature gradient in Washington State, USA. Xylem air-entry points were less negative in trees from moist environments (-0.71 and -1.32 MPa in the Hoh and Nisqually populations, respectively) than in trees from dry environments (-1.55 and -1.67 MPa in the Palouse and Yakima populations, respectively). Xylem cavitation in response to experimental drought was consistent with air-injection measures of xylem air-entry points for a given population. Populations vulnerable to cavitation also exhibited higher stem specific hydraulic conductivities and limited stomatal control compared with resistant populations. Populations exhibiting vulnerability to cavitation and limited stomatal control desiccated more rapidly during drought compared with resistant populations. This study provides evidence of interpopulation variation in resistance to drought-induced xylem cavitation, stomatal behavior, and hydraulic conductivity within Populus trichocarpa.     

Insights into xylem vulnerability to cavitation in Fagus sylvatica L.: phenotypic and environmental sources of variability

Xylem vulnerability to cavitation is a key parameter in understanding drought resistance of trees. We determined the xylem water pressure causing 50% loss of hydraulic conductivity (P(50)), a proxy of vulnerability to cavitation, and we evaluated the variability of this trait at tree and population levels for Fagus sylvatica. We checked for the effects of light on vulnerability to cavitation of stem segments together with a time series variation of P(50). Full sunlight-exposed stem segments were less vulnerable to cavitation than shade-exposed ones. We found no clear seasonal change of P(50), suggesting that this trait was designed for a restricted period. P(50) varied for populations settled along a latitudinal gradient, but not for those sampled along an altitudinal gradient. Moreover, mountainside exposure seemed to play a major role in the vulnerability to cavitation of beech populations, as we observed the differences along north-facing sides but not on south-facing sides. Unexpectedly, both north-facing mountainside and northern populations appeared less vulnerable than those grown on the southern mountainside or in the South of France. These results on beech populations were discussed with respect to the results at within-tree level.     

Water storage capacitance and xylem tension in isolated branches of temperate and tropical trees

Trees of tropical semi-deciduous forests range from "drought-avoiding" stem-succulent species with low-density wood (< 0.5 g cm(-3)), which maintain high stem water potentials (psi(STEM) > -0.7 MPa) throughout the year, to "drought-tolerant" deciduous hardwood species (wood density > 0.75 g cm(-3)), which dehydrate strongly during seasonal drought (psi(STEM) < -6 MPa). In stem-succulent and other drought-avoiding species, xylem vessels are surrounded by extensive parenchyma providing intracellular water storage, whereas in deciduous species stem water storage is mainly extracellular. Thirteen tropical and two temperate tree species, representing different functional types, were studied. The contribution of stem water storage to these species' water use during water stress was determined by time-series analysis of dehydration and rehydration of excised leaf-bearing branches of these trees. During dehydration, stem water potential slowly declined 1-2 MPa in drought-avoiding species, but in deciduous species it rapidly fell 4-5 MPa, suggesting that water storage capacitance was related to xylem anatomy. After immersion of dehydrated, leafless branches in water, the decline in xylem tension and rate of water uptake during rehydration were linearly related, as predicted by application of Ohm's law to water flux. The decline of xylem tension during rehydration was biphasic, with a phase of rapid water uptake into extracellular spaces being followed by a prolonged phase of slow water uptake into living cells. The rate of water uptake during rehydration and the minima of leaf water potential observed in the field during the dry season were highly correlated with water storage capacitance, indicating that wood anatomy is a major determinant of drought adaptation.     

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.     

Xylem cavitation in roots and stems of Douglas-fir and white fir

Roots of hardwoods have been shown to be more vulnerable to xylem cavitation than stems. This study examined whether this pattern is also observed in a conifer species. Vulnerability to cavitation was determined from the pressure required to inject air into the vascular system of hydrated roots and stems, and reduce hydraulic conductance of the xylem. According to the air-seeding hypothesis for the cavitation mechanism, these air pressures predict the negative xylem pressure causing cavitation in dehydrating stems. This was evaluated for stems of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and white fir (Abies concolor (Gord. & Glend.) Lindl.). The air-injection method was applied to roots and stems of different sizes and positions in Douglas-fir trees. Roots, especially smaller roots with a xylem diameter < 5 mm, were more vulnerable to cavitation than stems. Mean cavitation pressure for smaller roots was -2.09 +/- 0.42 versus -3.80 +/- 0.19 MPa for larger roots (> 8 mm diameter). Within the shoot system, smaller stems (< 5 mm diameter) were most vulnerable to cavitation, having a mean cavitation pressure of -4.23 +/- 0.565 versus -5.27 +/- 0.513 MPa for large stems (> 8 mm diameter). There was no correlation between tracheid diameter and mean cavitation pressure within root or stem systems, despite larger tracheid diameters in roots (23.3 +/- 3.9 micro m) than in stems (9.2 +/- 1.6 micro m). Smaller safety margins from cavitation in roots may be beneficial in limiting water use during mild drought, and in protecting the stem from low xylem pressures during extreme drought.     

Xylem vulnerability to cavitation in Pseudotsuga menziesii and Pinus ponderosa from contrasting habitats

In the Rocky Mountains, ponderosa pine (Pinus ponderosa (ssp.) ponderosa Dougl. ex P. Laws. & C. Laws) often co-occurs with Douglas-fir (Pseudotsuga menziesii var. glauca (Mayr) Franco). Despite previous reports showing higher shoot vulnerability to water-stress-induced cavitation in ponderosa pine, this species extends into drier habitats than Douglas-fir. We examined: (1) whether roots and shoots of ponderosa pine in riparian and slope habitats are more vulnerable to water-stress-induced cavitation than those of Douglas-fir; (2) whether species-specific differences in vulnerability translate into differences in specific conductivity in the field; and (3) whether the ability of ponderosa pine to extend into drier sites is a result of (a) greater plasticity in hydraulic properties or (b) functional or structural adjustments. Roots and shoots of ponderosa pine were significantly more vulnerable to water-stress-induced cavitation (overall mean cavitation pressure, Psi(50%) +/- SE = -3.11 +/- 0.32 MPa for shoots and -0.99 +/- 0.16 MPa for roots) than those of Douglas-fir (Psi(50%) +/- SE = -4.83 +/- 0.40 MPa for shoots and -2.12 +/- 0.35 MPa for roots). However, shoot specific conductivity did not differ between species in the field. For both species, roots were more vulnerable to cavitation than shoots. Overall, changes in vulnerability from riparian to slope habitats were small for both species. Greater declines in stomatal conductance as the summer proceeded, combined with higher allocation to sapwood and greater sapwood water storage, appeared to contribute to the ability of ponderosa pine to thrive in dry habitats despite relatively high vulnerability to water-stress-induced cavitation.     

Hydraulic differences along the water transport system of South American Nothofagus species: do leaves protect the stem functionality?

Hydraulic traits were studied for six Nothofagus species from South America (Argentina and Chile), and for three of these species two populations were studied. The main goal was to determine if properties of the water conductive pathway in stems and leaves are functionally coordinated and to assess if leaves are more vulnerable to cavitation than stems, consistent with the theory of hydraulic segmentation along the vascular system of trees in ecosystems subject to seasonal drought. Vulnerability to cavitation, hydraulic conductivity of stems and leaves, leaf water potential, wood density and leaf water relations were examined. Large variations in vulnerability to cavitation of stems and leaves were observed across populations and species, but leaves were consistently more vulnerable than stems. Water potential at 50% loss of maximum hydraulic efficiency (P(50)) ranged from -0.94 to -2.44 MPa in leaves and from -2.6 to -5.3 MPa in stems across species and populations. Populations in the driest sites had sapwood and leaves more vulnerable to cavitation than those grown in the wettest sites. Stronger diurnal down-regulation in leaf hydraulic conductance compared with stem hydraulic conductivity apparently has the function to slow down potential water loss in stems and protect stem hydraulics from cavitation. Species-specific differences in wood density and leaf hydraulic conductance (K(Leaf)) were observed. Both traits were functionally related: species with higher wood density had lower K(Leaf). Other stem and leaf hydraulic traits were functionally coordinated, resulting in Nothofagus species with an efficient delivery of water to the leaves. The integrity of the more expensive woody portion of the water transport pathway can thus be maintained at the expense of the replaceable portion (leaves) of the stem-leaf continuum under prolonged drought. Compensatory adjustments between hydraulic traits may help to decrease the rate of embolism formation in the trees more vulnerable to cavitation.     

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.     

Vulnerability of several conifers to air embolism

Hydraulic properties of xylem in seven species of conifer were studied during late winter and early spring 1991. Vulnerability to cavitation and air embolism was investigated using hydraulic conductivity and acoustic techniques. Embolisms were induced in branches excised from mature trees by air-drying them in the laboratory. Both techniques gave comparable results indicating that they both assess the same phenomenon. Within a tree, vulnerability was related to the permeability of the xylem, the largest stems tended to cavitate before the smallest ones when water deficits developed in a branch. Interspecific comparisons showed large differences in the xylem water potential needed to induce significant embolism, values ranged from -2.5 MPa in Pinus sylvestris to -4 MPa in Cedrus atlantica, but these differences did not correlate with differences in the xylem permeability of the species. The vulnerability of a species to air embolism was found to be consistent with its ecophysiological behavior in the presence of water stress, drought-tolerant species being less vulnerable than drought-avoiding species.     

Seasonal patterns of leaf gas exchange and water relations in dry rain forest trees of contrasting leaf phenology

Diurnal and seasonal patterns of leaf gas exchange and water relations were examined in tree species of contrasting leaf phenology growing in a seasonally dry tropical rain forest in north-eastern Australia. Two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., and two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret. were studied. The deciduous species had higher specific leaf areas and maximum photosynthetic rates per leaf dry mass in the wet season than the evergreens. During the transition from wet season to dry season, total canopy area was reduced by 70-90% in the deciduous species and stomatal conductance (g(s)) and assimilation rate (A) were markedly lower in the remaining leaves. Deciduous species maintained daytime leaf water potentials (Psi(L)) at close to or above wet season values by a combination of stomatal regulation and reduction in leaf area. Thus, the timing of leaf drop in deciduous species was not associated with large negative values of daytime Psi(L) (greater than -1.6 MPa) or predawn Psi(L) (greater than -1.0 MPa). The deciduous species appeared sensitive to small perturbations in soil and leaf water status that signalled the onset of drought. The evergreen species were less sensitive to the onset of drought and g(s) values were not significantly lower during the transitional period. In the dry season, the evergreen species maintained their canopies despite increasing water-stress; however, unlike Eucalyptus species from northern Australian savannas, A and g(s) were significantly lower than wet season values.     

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.     

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.     

Contrasting seasonal leaf habits of canopy trees between tropical dry-deciduous and evergreen forests in Thailand

We compared differences in leaf properties, leaf gas exchange and photochemical properties between drought-deciduous and evergreen trees in tropical dry forests, where soil nutrients differed but rainfall was similar. Three canopy trees (Shorea siamensis Miq., Xylia xylocarpa (Roxb.) W. Theob. and Vitex peduncularis Wall. ex Schauer) in a drought-deciduous forest and a canopy tree (Hopea ferrea Lanessan) in an evergreen forest were selected. Soil nutrient availability is lower in the evergreen forest than in the deciduous forest. Compared with the evergreen tree, the deciduous trees had shorter leaf life spans, lower leaf masses per area, higher leaf mass-based nitrogen (N) contents, higher leaf mass-based photosynthetic rates (mass-based P(n)), higher leaf N-based P(n), higher daily maximum stomatal conductance (g(s)) and wider conduits in wood xylem. Mass-based P(n) decreased from the wet to the dry season for all species. Following onset of the dry season, daily maximum g(s) and sensitivity of g(s) to leaf-to-air vapor pressure deficit remained relatively unchanged in the deciduous trees, whereas both properties decreased in the evergreen tree during the dry season. Photochemical capacity and non-photochemical quenching (NPQ) of photosystem II (PSII) also remained relatively unchanged in the deciduous trees even after the onset of the dry season. In contrast, photochemical capacity decreased and NPQ increased in the evergreen tree during the dry season, indicating that the leaves coped with prolonged drought by down-regulating PSII. Thus, the drought-avoidant deciduous species were characterized by high N allocation for leaf carbon assimilation, high water use and photoinhibition avoidance, whereas the drought-tolerant evergreen was characterized by low N allocation for leaf carbon assimilation, conservative water use and photoinhibition tolerance.     

Differential summer water use by Pinus edulis and Juniperus osteosperma reflects contrasting hydraulic characteristics

Previous studies of pinyon-juniper woodlands show that Pinus edulis Engelm. makes better use of soil water from summer precipitation pulses than does co-occurring Juniperus osteosperma (Torr.) Little. To investigate the basis of this difference, we examined seasonal variation in cavitation and hydraulic conductance. Pinus edulis remained isohydric over the growing season. Minimum water potentials never fell below -2.3 MPa, and the extent of xylem cavitation remained near constant during the dry season. In contrast, J. osteosperma was anisohydric, reaching water potentials as low as -6.9 MPa, and experiencing progressively greater xylem cavitation as the dry season progressed despite having more cavitation-resistant xylem than P. edulis. We conducted an irrigation experiment to observe the responses of the study species to a summer pulse of water. Although sap flow increased in both species in response to the 25-mm irrigation pulse, only J. osteosperma responded to the 10-mm pulse. This was inconsistent with the response of P. edulis to light rain events and may have been due to a difference in the distribution of irrigation water and rain water between the under- and between-canopy areas. Whole-plant conductance increased following the 25-mm irrigation in P. edulis but remained constant in J. osteosperma. We hypothesized that this difference was caused, in part, by differential refilling of embolized xylem. Area specific hydraulic conductivity was 66% higher in roots of irrigated P. edulis trees relative to roots of control trees 3 days after the 25-mm irrigation (t = 2.14, P = 0.02, df = 16). There was no change in hydraulic conductivity of the roots of J. osteosperma or in the stems of either species. Our results indicate that the response to an irrigation pulse in P. edulis depended on cavitation avoidance in stems and the reversal of cavitation in roots, resulting in increased whole-plant conductance and water uptake. In contrast, J. osteosperma failed to exploit light summer rain events but was able to extract deep soil water at low water potentials.     

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.     

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).     

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.