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Bamboos are arborescent monocotyledons that have no secondary growth, but can continually produce conduits with diameters appropriate to the current size of the plant. Here, we studied bamboo hydraulic architecture to address the mechanisms involved in compensating for the increase in hydraulic resistance during ontogeny. We measured the hydraulic weighted vessel diameters (Dh ) at different distances from the apex along the stem of Bambusa textilis. The hydraulic resistance of different components and individuals of different heights were quantified using the high-pressure flowmeter method. The Dh showed tip-to-base widening with a scaling exponent in the range of those reported for trees. Although theoretical hydraulic conductivity decreased from base-to-tip, leaf-specific conductivity did not change. Leaves contributed the most to the whole-shoot hydraulic resistance, followed by the leaf-bearing branches. Roots contributed c. 13% to whole-plant resistance. Interestingly, taller individuals showed lower whole-shoot resistance owing to an increased number of resistances in parallel (side-branches), while leaf-specific resistance was independent of plant size. Tip-to-base vessel widening and height-independent constant leaf-specific conductance could be mechanisms for hydraulic optimization in B. textilis. Similar patterns have also been found in woody plants with secondary growth, but this bamboo exhibits them without secondary growth.

Xylem vulnerability to embolism represents an important trait to determine species distribution patterns and drought resistance. However, estimating embolism resistance frequently requires time-consuming and ambiguous hydraulic lab measurements. Based on a recently developed pneumatic method, we present and test the "Pneumatron", a device that generates high time-resolution and fully automated vulnerability curves. Embolism resistance is estimated by applying a partial vacuum to extract air from an excised xylem sample, while monitoring the pressure change over time. Although the amount of gas extracted is strongly correlated with the percentage loss of xylem conductivity, validation of the Pneumatron was performed by comparison with the optical method for Eucalyptus camaldulensis leaves. The Pneumatron improved the precision of the pneumatic method considerably, facilitating the detection of small differences in the (percentage of air discharged [PAD] < 0.47%). Hence, the Pneumatron can directly measure the 50% PAD without any fitting of vulnerability curves. PAD and embolism frequency based on the optical method were strongly correlated (r2 = 0.93) for E. camaldulensis. By providing an open source platform, the Pneumatron represents an easy, low-cost, and powerful tool for field measurements, which can significantly improve our understanding of plant-water relations and the mechanisms behind embolism.

Cavitation resistance is a key trait for characterizing the drought adaption in plants and is usually presented in terms of vulnerability curves. Three principal techniques have been developed to produce vulnerability curves, but curves generated with centrifugation are reported to suffer from artifacts when applied to long-vesseled species. The main cause of this artifact is the issue of open vessels, resulting in a nano-particle effect that may seed premature embolism. We used two methods to test the potential mechanism behind the nano-particle effect in centrifuge-based vulnerability curves. A four-cuvette rotor system based on a traditional Cochard rotor was designed to inhibit effervescence while injecting water, but the recalcitrant vulnerability curves in Robinia could not be eliminated. There may be multiple sources, besides effervescence, of hypothetical nano-particles: they may arise from cut surfaces or they may be always present in the injected water, leading to the premature embolisms. To prevent the entry of the hypothetical nano-particles, water extraction curves in terms of PLV (percentage loss volume of extracted water from stems) vs tensions were constructed. The PLV curves of Robinia showed s-shaped characteristics after subtracting the first Weibull components from water extraction curves, which were not related to the water loss from vessels according to dye staining experiments. The differences between T50 (xylem tension at which 50% of hydraulic conductivity is lost) in mean PLV curve and T50 in percentage loss of conductivity curves determined by the four-cuvette rotor system and by the bench dehydration method were 3.9 MPa and 0.7 MPa, respectively. Hence, PLV curves may be a valid way to measure the cavitation resistance in long-vesseled species with centrifugation. Keeping bark intact in the process of measurement is recommended, otherwise it would increase evaporation from the entire system.

Dwarfing of fruit trees is often achieved through the use of dwarfing rootstocks. Dwarf trees are characterized by sustained reductions in vegetative growth during the lifetime of the tree. The dwarfing mechanism is not well understood, but it has been hypothesized that hydraulic properties of the rootstock and the graft union are involved. It is hypothesized here that leaf- or stem-specific resistance of at least one hydraulic component of the water transport system would be negatively correlated with rootstock 'vigour', and this could be useful for selection of rootstocks. Hydraulic resistance (R) of fully grown apple trees on a variety of rootstocks of different 'vigours' was measured. Most measurements were with the evaporative flux (EF) method, where water uptake measured with sap flow sensors was related to the pressure gradient from soil (taken as pre-dawn leaf) and midday root (taken as covered root-sucker), stem (from covered leaf), and exposed and shaded leaf water potentials (Psi(l)). R of trees on dwarfing M9 rootstock was compared with that of more vigorous MM106 and MM111 rootstocks in Israel and Vermont, USA. In Israel, M9 consistently had higher leaf-specific hydraulic resistance (R(l)) in the soil to scion stem pathway, but this difference was only significant for one summer. R was larger in M9 between the root and stem, implicating the graft union as the site of increased resistance. In Vermont, R(l) of 9- and 10-year-old trees on six rootstocks of various vigours was not consistently related to vigour, and stem-specific resistance (R(s)) increased with increasing vigour. High pressure flow meter (HPFM) measurements gave a lower R than the EF method in all but one case, perhaps indicating a significant amount of xylem dysfunction in these trees, and demonstrated the increased resistivity of stem sections that included dwarf graft unions as compared with non-graft stem sections. It is concluded that stem- and leaf-specific R are not consistently positively correlated with dwarfing, although the increased resistivity of the graft union in dwarfing rootstocks may influence the transport of water and other elements across the graft union, and therefore be involved in the dwarfing mechanism.

Cavitation of water in xylem vessels followed by embolism formation has been authenticated for more than 40 years. Embolism formation involves the gradual buildup of bubble pressure (air) to atmospheric pressure as demanded by Henry's law of equilibrium between gaseous and liquid phases. However, the tempo of pressure increase has not been quantified. In this report, we show that the rate of pressurization of embolized vessels is controlled by both fast and slow kinetics, where both tempos are controlled by diffusion but over different spatial scales. The fast tempo involves a localized diffusion from endogenous sources: over a distance of about 0.05 mm from water-filled wood to the nearest embolized vessels; this process, in theory, should take2 min. The slow tempo involves diffusion of air from exogenous sources (outside the stem). The latter diffusion process is slower because of the increased distance of diffusion of up to 4 mm. Radial diffusion models and experimental measurements both confirm that the average time constant is17 h, with complete equilibrium requiring 1 to 2 d. The implications of these timescales for the standard methods of measuring percentage loss of hydraulic conductivity are discussed in theory and deserve more research in future.

Summary • The leaf hydraulic conductance (Kleaf) is a major determinant of plant water transport capacity. Here, we measured Kleaf, and its basis in the resistances of leaf components, for fully illuminated leaves of five tree species that regenerate in deep shade, and five that regenerate in gaps or clearings, in Panamanian lowland tropical rainforest. We also determined coordination with stomatal characters and leaf mass per area. • Kleaf varied 10-fold across species, and was 3-fold higher in sun- than in shade-establishing species. On average, 12% of leaf hydraulic resistance (= 1/Kleaf) was located in the petiole, 25% in the major veins, 25% in the minor veins, and 39% outside the xylem. Sun-establishing species had a higher proportion of leaf resistance in the xylem. Across species, component resistances correlated linearly with total leaf resistance. • Kleaf correlated tightly with indices of stomatal pore area, indicating a coordination of liquid- and vapor-phase conductances shifted relative to that of temperate woody species. • Leaf hydraulic properties are integrally linked in the complex of traits that define differences in water use and carbon economy across habitats and vegetation zones.

This paper reports how water stress correlates with changes in hydraulic conductivity of stems, leaf midrib, and whole leaves ofPrunus laurocerasus. Water stress caused cavitation-induced dysfunction in vessels of P. laurocerasus. Cavitation was detected acoustically by counts of ultrasonic acoustic emissions and by the loss of hydraulic conductivity measured by a vacuum chamber method. Stems and midribs were approximately equally vulnerable to cavitations. Although midribs suffered a 70% loss of hydraulic conductance at leaf water potentials of −1.5 MPa, there was less than a 10% loss of hydraulic conductance in whole leaves. Cutting and sealing the midrib 20 mm from the leaf base caused only a 30% loss of conduction of the whole leaf. A high-pressure flow meter was used to measure conductance of whole leaves and as the leaf was progressively cut back from tip to base. These data were fitted to a model of hydraulic conductance of leaves that explained the above results, i.e. redundancy in hydraulic pathways whereby water can flow around embolized regions in the leaf, makes whole leaves relatively insensitive to significant changes in conductance of the midrib. The onset of cavitation events in P. laurocerasus leaves correlated with the onset of stomatal closure as found recently in studies of other species in our laboratory.

Recovery of hydraulic conductivity after the induction of embolisms was studied in woody stems of laurel (Laurus nobilis). Previous experiments confirming the recovery of hydraulic conductivity when xylem pressure potential was less than −1 MPa were repeated, and new experiments were done to investigate the changes in solute composition in xylem vessels during refilling. Xylem sap collected by perfusion of excised stem segments showed elevated levels of several ions during refilling. Stem segments were frozen in liquid N2 to view refilling vessels using cryoscanning electron microscopy. Vessels could be found in all three states of presumed refilling: (a) mostly water with a little air, (b) mostly air with a little water, or (c) water droplets extruding from vessel pits adjacent to living cells. Radiographic probe microanalysis of refilling vessels revealed nondetectable levels of dissolved solutes. Results are discussed in terms of proposed mechanisms of refilling in vessels while surrounding vessels were at a xylem pressure potential of less than −1 MPa. We have concluded that none of the existing paradigms explains the results.

Quasi-steady-state measurements of root hydraulic conductance (KR) of Olea oleaster Hoffmgg. et Link potted seedlings were performed using a pressure chamber with the aim of: (a) measuring the impact of different water-stress levels on a KR; (b) measuring the kinetics of KR recovery several days after soil rewetting; (c) relating changes in KR to changes in root anatomy and morphology. Increasing water-stress was applied in terms of ratio of leaf water potential (ΨL) measured at midday to that at zero turgor (ΨTLP), i.e. ΨL/ΨTLP=0·5, 1·0, 1·2, 1·6; KR was measured initially and at 24, 48, 72, 96 h after irrigation. Values of KR in seedlings stressed to ΨL/ΨTLP=1·2 increased for 48 h after irrigation from 0·23 to 0·97×10−5 kg s−1 m−2 MPa−1 i.e. from 16% to 66% of that measured in unstressed seedlings. A marked shift of the x-axis intercept of the straight line relating flow to pressure (zero flow at non-zero pressure) was recorded initially after irrigation and persisted up to 48 h. Recovery of KR occurred within 24 h after irrigation in seedlings at ΨL/ΨTLP=0·5 and 48 h later in those at ΨL/ΨTLP=1·0. Severe drought stress (ΨL/ΨTLP=1·6) caused anatomical changes to roots which formed a two-layered exodermis with thicker suberized walls and a three- to four-layered endodermis with completely suberized tangential walls. Recovery of KR in these roots required resumed growth of root tips and emergence of new lateral roots.

Occasional droughts may be important in controlling the distribution and structure of forest types in relatively aseasonal north Borneo. The low water retention capacity of the coarse, sandy soils on which tropical heath forest occurs may cause drought to develop more quickly and severely than on the finer textured soils of nearby dipterocarp forest. Resistance to drought-induced embolism is considered an important component of drought tolerance. We constructed embolism vulnerability curves relating loss in hydraulic conductivity to xylem tension by the air-injection method for understory trees of 14 species from both tropical heath and mixed dipterocarp forests in Brunei Darussalam. There was no significant difference (Mann-Whitney U-test, P = 0.11) between forest types in the xylem tension at which 50% loss of hydraulic conductivity occurred. Most species from both forest types were highly vulnerable to embolism compared with species from seasonal tropical forests. We speculate that other mechanisms, such as stomatal control to prevent development of embolism-inducing xylem tensions, are more cost-effective adaptations against occasional drought, but that the attendant reduction in productivity and competitive ability places a greater premium on resistance to embolism when drought is annual and predictable.

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.

The functional xylem anatomy and the hydraulic conductivity of intact and treated branch junctions of the diffuse-porous sugar maple (Acer saccharum Marsh.) were compared to those of the ring-porous black oak (Quercus velutina Lam.). Maple shoots possessed greater growth intensity than those of oak. The extensive growth of the maple trees resulted in about a two-fold increase in xylem production in the maple branches. Branches were altered by removing a patch of bark from the base of a branch (near a junction) leaving a bridge of bark on the upper or lower side of the branch. The experimentally treated branch junctions revealed that, in oak, most (up to 92%) of the water flows in the lower side of a branch, where most of the large vessels occurred. In maple, most of the conductive tissue was observed to form in the upper side of the branches, which was equally or more conductive than the lower side. A treatment of longitudinal, parallel scratches in the bark-bridge, which reduced earlywood vessel width, substantially decreased conductivity (to only 15%) in oak, but had no effect on conductivity in maple. In maple, such wounding stimulated more wood formation and increased conductivity. In both trees, a narrow bridge at the junction induced more wood formation and higher conductivity in the branch. The mechanisms controlling wood formation and water flow in branch junctions of ring- and diffuse-porous trees are discussed.

Several parameters related to the water relations of eight woody hemiepiphytes with different photosynthetic pathways were studied in situ and in the laboratory on Barro Colorado Island, Panama. As a group, woody hemiepiphytes tended to have less conductive stems per unit leaf area (lower k L ) and invested less wood cross-section per unit leaf area compared to free-standing trees, while their specific conductivity (K s ) was significantly higher. Among hemiepiphytes, there were significant differences between C 3 and CAM (Crassulacean Acid Metabolism) species in respect to leaf characteristics, transpiration rates, diel patterns of water flow through aerial roots, and in hydraulic architecture parameters. Average transpiration rates of the two Clusia species (C 3 -CAM) were lower by about an order of magnitude compared to the C 3 species. In all C 3 species, sap flow through aerial roots (F) closely followed transpiration (E), whereas E and F were decoupled in time in the C 3 -CAM species Clusia uvitana: considerable long-distance water flow occurred at night. The hydraulic efficiency of the other C 3 -CAM species, C. minor, was the lowest of the five species investigated.

Pressure probe measurements have been interpreted as showing that xylem pressures below c. -0.4 MPa do not exist and that pressure chamber measurements of lower negative pressures are invalid. We present new evidence supporting the pressure chamber technique and the existence of xylem pressures well below -0.4 MPa. We deduced xylem pressures in water-stressed stem xylem from the following experiment : (1) loss of hydraulic conductivity in hydrated stem xylem (xylem pressure = atmospheric pressure) was induced by forcing compressed air into intact xylem conduits ; (2) loss of hydraulic conductivity from cavitation and embolism in dehydrating stems was measured, and (3) the xylem pressure in dehydrated stems was deduced as being equal and opposite to the air pressure causing the same loss of hydraulic conductivity in hydrated stems. Pressures determined in this way are only valid if cavitation was caused by air entering the xylem conduits (air-seeding). Deduced xylem pressure showed a one-to-one correspondence with pressure chamber measurements for 12 species (woody angiosperms and gymnosperms) ; data extended to c. -10 MPa. The same correspondence was obtained under field conditions in Betula occidentalis Hook., where pressure differences between air- and water-filled conduits were induced by a combination of in situ xylem water pressure and applied positive air pressure. It is difficult to explain these results if xylem pressures were above -0.4 MPa, if the pressure chamber was inaccurate, and if cavitation occurred by some mechanism other than air-seeding. A probable reason why the pressure probe does not register large negative pressures is that, just as cavitation within the probe limits its calibration to pressures above c. -0.5 MPa, cavitation limits its measurement range in situ.

Summary • It is widely assumed that post-fire tree mortality results from necrosis of phloem and vascular cambium in stems, despite strong evidence that reduced xylem conductivity also plays an important role. • In this study, experiments with Populus balsamifera were used to demonstrate two mechanisms by which heat reduces the hydraulic conductivity of xylem: air seed cavitation and conduit wall deformation. Heat effects on air seed cavitation were quantified using air injection experiments that isolate potential temperature-dependent changes in sap surface tension and pit membrane pore diameters. Heat effects on conduit wall structure were demonstrated using air conductivity measurements and light microscopy. • Heating increased vulnerability to cavitation because sap surface tension varies inversely with temperature. Heating did not affect cavitation via changes in pit membrane pore diameters, but did cause significant reductions in xylem air conductivity that were associated with deformation of conduit walls (probably resulting from thermal softening of viscoelastic cell wall polymers). • Additional work is required to understand the relative roles of cavitation and deformation in the reduction of xylem conductivity, and how reduced xylem conductivity in roots, stems, and branches correlates and interacts with foliage and root necroses to cause tree mortality. Future research should also examine how heat necrosis of ray parenchyma cells affects refilling of embolisms that occur during and after the fire event.

SUMMARY Clusia uvitana Pittier (Clusiacea) is a tropical hemiepiphyte that has been shown to display a high plasticity in the expression of CAM in response to the environment. When water is available CO2 is taken up mostly during the- day. This study of the water relations and hydraulic architecture has revealed that leaf water potentials, £ ranged from 0-7 to -0.9 MPa and changed very little with time or water availability. The absolute hydraulic conductivity of stem segments (K,) and the specific conductivity (K1) were comparable to many other temperate and tropical species, but the leaf specificity conductivity (K1) was 1/3 to 1/30 that of many other species. So stems supported high leaf areas per unit of hydraulic conductivity. C uvitana was very vulnerable to cavitation, reaching 50 % loss of hydraulic conductivity at stem £=1.3 MPa. The species survives in spite of low K1 and high xylem vulnerability, because the CAM physiology insures low transpiration rates and high ability to evade dehydration.

The occurrence of root pressure, the vulnerability of xylem vessels to drought-induced cavitation, and the seasonal changes in hydraulic conductivity due to embolism were studied in the culms of Rhipidocladum racemiflorum (Steud.) McClure, a tropical vine-like bamboo from central Panama. Positive hydrostatic potentials up to 120 kPa occurred only during the wet season when the transpiration rate of the plant was low, i.e. at night or during rain events. Although the xylem vessels were large and efficient for conducting water, they were highly resistant to cavitation. Xylem water potentials lower than -4.5 MPa were required to induce 50% loss of hydraulic conductivity in culms

Hydraulic resistance to water flow was measured in the xylem and leaves of above-ground portions of Acer saccharum and Acer rubrum for trees with trunk diameters (D) ranging from 0.02 to 0.2 m. Resistance (area basis) to water flow in leaves and petioles was 24 and 13×10 3 MPa m 2 s kg -1 for A. saccharum and A. rubrum, respectively. Leaf area of whole trees was proportional to D 1.82 . Absolute xylem resistance (MPa s kg -1 ) was proportional to D -1.64 . So xylem resistance (area basis) was proportional to D 0.2 , which was not significantly different from a zero dependence (D 0 ) for D≤0.2 m

Vulnerability curves (VCs) measure the ability of vessels to retain metastable water without embolisms that lower the hydraulic conductivity of stems. The fastest method of measuring VCs is the centrifuge technique and the Cochard cavitron is a method that allows measurement of hydraulic conductivity of stems while they are spinning. This paper describes the pattern of embolism that results after spinning the stems of hybrid aspen (Populus tremula×P. tremuloides) and two hybrid cottonwoods (P38P38 P. balsamifera×P. simonii and Northwest, which is a hybrid of P. deltoides×P. balsamifera). It is recognized that the pattern of embolism induced in a centrifuge ought to differ from the pattern during natural dehydration of plants because the profiles of tension vs distance greatly differ under the two modes of inducing stress. The pattern of embolism was visualized by a staining technique and quantified by traditional measurements of percentage loss conductivity (PLC) performed on subsample segments excised from spun stems. We found a pattern of embolism approximating that expected from theory: (1) PLC near the axis of rotation exceeded the average; (2) PLC was quite high near the ends of the stems, even though tension ought to be zero; (3) large vessels cavitated before small vessels; (4) more embolism occurred near the base than near the apex of the stems. However, we could not always scale up from subsample conductivity and PLC to whole-stem conductivity. This pattern of embolism is interpreted in terms of vessel diameter and vessel length.

The objective of this study was to quantify the relationship between vulnerability to cavitation and vessel diameter within a species. We measured vulnerability curves (VCs: percentage loss hydraulic conductivity versus tension) in aspen stems and measured vessel-size distributions. Measurements were done on seed-grown, 4-month-old aspen (Populus tremuloides Michx) grown in a greenhouse. VCs of stem segments were measured using a centrifuge technique and by a staining technique that allowed a VC to be constructed based on vessel diameter size-classes (D). Vessel-based VCs were also fitted to Weibull cumulative distribution functions (CDF), which provided best-fit values of Weibull CDF constants (c and b) and P(50) = the tension causing 50% loss of hydraulic conductivity. We show that P(50) = 6.166D(-0.3134) (R(2) = 0.995) and that b and 1/c are both linear functions of D with R(2)0.95. The results are discussed in terms of models of VCs based on vessel D size-classes and in terms of concepts such as the 'pit area hypothesis' and vessel pathway redundancy.

Experiments were conducted to determine the influence of stem diameter, xylem pressure potential, and temperature on the rate of recovery of hydraulic conductivity in embolized stems of Acer saccharum Marsh. Recovery of conductivity was accompanied by an increase in stem water content as water replaced air bubbles and bubbles dissolved from vessels into the surrounding water. The time required for stems to go from less than 3 to 100% hydraulic conductivity increased approximately with the square of the stem diameter and increased with decreasing xylem pressure potential. Recovery was halted when xylem pressure potential decreased below -6 kPa. Increasing xylem pressure from 13 to 150 kPa reduced the time for recovery by a factor of 4. Temperature had little influence on the rate of recovery of hydraulic conductivity. All of these results are in accord with a theory of bubble dissolution in which it is assumed that: (a) the rate of bubble dissolution is rate limited by diffusion of air from the bubbles to the outer surface of the stems, (b) the equilibrium concentration of gases in liquid in stems is determined by Henry's law at all air-water interfaces, (c) the equilibrium solubility concentration is determined only by the partial pressure of the gas in the gas phase and not directly by the liquid-phase pressure, and (d) the gas pressure of an entrapped air bubble in the lumen of a cell can never be less than atmospheric pressure at equilibrium.

Loss of hydraulic conductivity occurs in stems when the water in xylem conduits is subjected to sufficiently negative pressure. According to the air-seeding hypothesis, this loss of conductivity occurs when air bubbles are sucked into water-filled conduits through micropores adjacent to air spaces in the stem. Results in this study showed that loss of hydraulic conductivity occurred in stem segments pressurized in a pressure chamber while the xylem water was under positive pressure. Vulnerability curves can be defined as a plot of percentage loss of hydraulic conductivity versus the pressure difference between xylem water and the outside air inducing the loss of conductivity. Vulnerability curves were similar whether loss of conductivity was induced by lowering the xylem water pressure or by raising the external air pressure. These results are consistent with the air-seeding hypothesis of how embolisms are nucleated, but not with the nucleation of embolisms at hydrophobic cracks because the latter requires negative xylem water pressure. The results also call into question some basic underlying assumptions used in the determination of components of tissue water potential using "pressure-volume" analysis.

A theoretical model of bubble dissolution in xylem conduits of stems was designed using the finite differential method and iterative calculations via computer. The model was based on Fick's, Henry's and Charles' laws and the capillary equation. The model predicted the tempo of recovery from embolism in small diameter branches of woody plants with various xylem structures under different xylem water pressures. The model predicted the time required to recover conductivity in any position in the stem. Repeated iterative solution of the model for different situations yielded an empirical formula to calculate the time for complete recovery of conductivity in stems from a fully embolised initial state. The time, tp, is given by: where α is a temperature coefficient; D is the coefficient of diffusion of air in wood at 25°C; rcs is the ratio of the area of total conduit cross section to the stem cross section; Ψxp is the stem xylem pressure potential (Pa, where 0 Pa equals atmospheric pressure); τ is solution surface tension (0.072 N m−1); and Dc and Ds are diameters of the conduits and the stem, respectively (m). The equation is valid only when Ψxp > –4τ/Dc. The model predicts no recovery of conductivity when Ψxp≤–4τ/Dc. The model agreed with experiments.

Summary 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.

The water relations and hydraulic architecture of a tropical tree (Schefflera morototoni) and of two temperate species (Acer saccharum and Thuja occidentalis) are reported. Among the water relations parameters measured were leaf and stem water storage capacity, leaf water potential, transpiration, and vulnerability of stems to cavitation and loss of hydraulic conductivity by embolisms. Among the hydraulic architecture parameters measured were hydraulic conductivity per unit pressure gradient, specific conductivity, leaf-specific conductivity, and Huber value. In terms of vulnerability of stems to cavitation, stem and leaf capacitances, and leaf-specific conductivity, all three species followed the same sequence: Schefflera > Acer > Thuja. It is argued here that the high stem capacitance and high leaf-specific conductivity of Schefflera are necessary to compensate for its high vulnerability to cavitation. Extractable water storage per unit leaf area in Schefflera stems is >100 times that of Acer and may permit the species to survive unusually long, dry seasons in Panama. Although Schefflera frequently grows >20 meters, the biggest resistance to water flow in the shoots resides in the leaves.

The mechanism of water-stress-induced xylem embolism was studied in three species of conifers: Abies balsamea (L.) Mill., Picca rubens Sarg, and Juniperus virginiana L. Each species showed a characteristic relationship between xylem tension and the loss of hydraulic conductivity by air embolism. Abics balsamea and Picca rubens began to embolize at tensions between 2 and 3 MPa and were completely non-conducting between 3 and 4 MPa. Juniperus virginiana was least vulnerable, beginning to embolize at 4 and still retaining approximately 10% conductivity at 10 MPa. As with a previous study of the vessel-bearing Accr saccharum Marsh., a brief perfusion of branch segments with an oxalic acid and calcium solution (10 and 0.1 mol m−3. respectively) increased the vulnerability of the xylem to embolism; this was especially pronounced in Abies balsamea. In order to test whether embolism was caused by aspiration of air into functional tracheids from neighbouring embolized, ones (the ‘air-seeding’hypothesis), hydrated branch segments were injected with air at various pressures and measured for embolism. Results supported the air-seeding hypothesis because the relationship between injection pressure and embolism for both native and oxalic-calcium-treated segments was essentially the same as for embolism induced by xylem tension. Structural and experimental evidence suggested the air seeding occurred through inter-tracheid pit membranes when the thickened torus region of the membrane became displaced from its normal sealing position over the pit aperture. Thus, the embolism-inducing tension may be a function of pit membrane flexibility. This tension is of ecological significance because it reflects to some extent the range of xylem tensions to which a species is adapted.

Bond's "slow seedling" hypothesis proposes that, because of slow growth rates caused by an inefficient transport system and low leaf photosynthetic capacity, gymnosperm seedlings are weak competitors with angiosperms in productive habitats. We measured component (shoot, leaf, and root) and whole-plant hydraulic conductances of sapling-sized tropical plants growing on nitrogen-poor white sand in Borneo. After accounting for size effects, there were no significant differences in conductances between evergreen angiosperms (nine species) and conifers (three species). Plant successional status or transpiration rate seemed more important than soil fertility in determining hydraulic conductance-colonizers had significantly higher whole-plant conductance than late-succession species. Contrary to prediction, leaf hydraulic conductance (normalized by projected leaf area) was unrelated to complexity of venation in conifers and angiosperms, but was highly correlated with whole-plant conductance. Analyses of published data showed that leafless branches of temperate deciduous angiosperms had higher leaf-area normalized hydraulic conductivity than conifers, but there was no significant difference in adult, whole-plant conductance between these taxa. Thus, at the branch level, conifers with narrow tracheids have less efficient transport than angiosperms with wider vessels, but variations in other resistance components and hydraulic architecture (e.g., sapwood/leaf area ratio) ultimately equalize the sufficiency of water transport to leaves of conifers and angiosperms. Although failing to support one of the proposed mechanisms, our findings did not refute the "slow seedling" hypothesis per se.

Studies of the desiccation tolerance of 15-month-old Licania platypus (Hemsl.) Fritsch seedlings were performed on potted plants. Pots were watered to field capacity and then dehydrated for 23-46 d to reach various visible wilting stages from slightly-wilted to dead. Root hydraulic conductance, k(r), was measured with a high-pressure flow meter and whole-stem hydraulic conductance, k(ws), was measured by a vacuum chamber method. Leaf punches were harvested for measurement of leaf water potential by a thermocouple psychrometer and for measurement of fresh- and dry-weight. L. platypus was surprisingly desiccation-tolerant, suggesting that most species of central Panama may be well adapted to the seasonality of rainfall in the region. The slightly-wilted stage corresponded to leaf water potentials and relative water contents of -2.7 MPa and 0.85, respectively, but plants did not die until these values fell to -7.5 MPa and 0.14, respectively. As desiccation proceeded k(r) and k(ws) declined relative to irrigated controls, but k(ws) was more sensitive to desiccation than k(r). Values of k(ws) declined by 70-85% in slightly-wilted to dead plants, respectively. By comparison, k(r) showed no significant change in slightly-wilted plants and fell by about 50% in plants having severely-wilted to dead shoots.

The molecular function of several aquaporins is well characterized, e.g. by analysis of aquaporin-expressing Xenopus oocytes. However, their significance in the physiology of water transport of multicellular organism is still uncertain in many cases. The tobacco plasma membrane aquaporin NtAQP1 was used to elucidate this issue. By comparing antisense plants which were inhibited in NtAQP1-expression to control plants. we found evidence for NtAQP1-function in cellular-and whole plant water-relations- Consequences of a decrease in cellular water permeability were determined by measurement of transpiration rate, stem and leaf water potential as well as growth experiments under extreme soil water depletion. Plants impaired in NtAQP1-expression showed a reduced root hydraulic conductivity and a lower water-stress resistance. In conclusion, the results give emphasis to the importance of a symplastic aquaporin-mediated water transport in whole plant water-relations.

An overview of the relationship between plant gas exchange, the potential hydraulic gradient, the size of the plant and its hydraulic conductance is presented. Key references are used to exemplify arguments of whole-plant optimality and to explain the origin and development of the dominant paradigm for interpreting the nature of water use and growth in plants.

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.

This chapter reviews how hydraulic architectures of tropical trees and lianas (woody vines) influence the flow of water from roots to leaves. The hydraulic design potentially can limit plant water relations, gas exchange, successional distribution, and even the maximum height (or length, in the case of lianas) that a species can attain. Important parameters include vulnerability to drought-induced cavitation (since embolism reduces hydraulic conductance), root pressures (since these potentially could result in re-filling of conduits following cavitation events, thereby increasing conductance), leaf specific conductivity (which, together with transpiration rates, can predict pressure gradi­ents throughout the plant), and water storage capacity (since this might determine the ability to survive water shortage). Some of the issues that are dealt with here are the impact of vessel diameter on drought- and freezing-induced embolism, the role of root pressures in the occasional removal of embolisms, and the ways in which the hydraulic architecture differs in different growth forms such as trees, shrubs, lianas, and hemiepiphytes. The ecological and physiological trade-offs of different architectures are discussed, and comparisons are made with temperate plants.

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.

Steady-state and dynamic methods were used to measure the conductivity to water flow in large woody root systems. The methods were destructive in that the root must be excised from the shoot but do not require removal of the root from the soil. The methods involve pushing water from the excised base of the root to the apex, causing flow in a direction opposite to that during normal transpiration. Sample data are given for two tropical (Cecropia obtusifolia and Lacistema aggregatum) and two temperate species (Acer saccharum and Juglans regia cv Lara). A hysteresis was observed in the relationship between applied pressure and resulting flow during dynamic measurements. A mathematical model (AMAIZED) was derived for the dynamics of solute and water flow in roots. The model was used to interpret results obtained from steady-state and dynamic measurements. AMAIZED is mathematically identical with the equations that describe Munch pressure flow of solute and water in the phloem of leaves. Results are discussed in terms of the predictions of AMAIZED, and suggestions for the improvement of methods are made.

Hydraulic conductivities were measured in branch junctions and in the proximal segments of Quercus velutina Lam., Acer saccharum Marsh., and Tsuga canadensis (L.) Carr. In all three species, the basal proximal segment was more conductive than the junction by a factor of 1.1–1.5. There was no consistent pattern for the distal proximal segments, where the conductivities were higher, lower, or the same as the junction. These data are discussed in terms of Martin H. Zimmermann's concept of plant segmentation. It is concluded that junction constrictions to water flow contribute less to plant segmentation than the variation in leaf specific conductivity in the crown of these species.

Trees of Juglans regia L. shed leaves when subjected to drought. Before shedding (when leaves are yellow), the petioles have lost 87% of their maximum hydraulic conductivity, but stems have lost only 14% of their conductivity. This is caused by the higher vulnerability of petioles than stems to water-stress induced cavitation. These data are discussed in the context of the plant segmentation hypothesis.

Plant water relations of nine woody species were studied in a lower montane rain forest in Panama. These data provide a partial test of the hypothesis that hydraulic architecture of lower montane species might limit transpiration and thus leaf size or nutrient transport (as suggested by J. Cavelier and E. G. Leigh, respectively). Diurnal variation in leaf transpiration was closely correlated with changes in net radiation. Peak transpiration rates (7 × 10–5 kg s–1 m–2) were as high as peak transpiration rates from tropical lowland forests but mean daily water use [0.39 ± 0.08 (SEM) kg m–2 day–1] were mostly lower than comparable data from tropical lowland forests. Thus transpiration rates are sufficiently high for sufficiently long periods to make it unlikely that nutrient transport is limited by transpiration. Another objective of this study was a comparison of two different methods to measure hydraulic conductance (Kh = flow rate per unit pressure gradient) and leaf specific conductance of stem segments (KL = Kh/leaf area distal to the segment). The results obtained with the traditional conductivity apparatus and the high pressure flow meter method, yielded similar results in six out of seven cases.

The dynamics of growth (shoot and root dry weights, surface areas, hydraulic conductances, and root length) were measured in seedlings of five neotropical tree species aged 4-16 months. The species studied included two light-demanding pioneers (Miconia argentea and Apeiba membranacea) and three shade-tolerant young- or old-forest species (Pouteria reticulata, Gustavia superba, and Trichilia tuberculata). Growth analysis revealed that shoot and root dry weights and hydraulic conductances and leaf area all increased exponentially with time. Alternative methods of scaling measured parameters to reveal differences that might explain adaptations to microsites are discussed. Scaling root conductance to root surface area or root length revealed a few species differences but nothing that correlated with adaptation to light regimes. Scaling of root surface area or root length to root dry weight revealed that pioneers produced significantly more root area and length per gram dry weight investment than shade-tolerant species. Scaling of root and shoot hydraulic conductances to leaf area and scaling of root conductance to root dry weight and shoot conductance to shoot dry weight also revealed that pioneers were significantly more conductive to water than shade-tolerant species. The advantages of scaling hydraulic parameters to leaf surface area are discussed in terms of the Ohm's law analogue of water flow in plants.

Hydraulic conductivity of the xylem is computed as the quotient of mass flow rate and pressure gradient. Measurements on excised plant stems can be difficult to interpret because of time-dependent reductions in flow rate, and because of variable degrees of embolism. Using Acer saccharum Marsh. stems, we found that certain perfusing solutions including dilute fixatives (e.g. 0.05% formaldehyde) and acids with pH below 3 (e.g. 10 mol m−3 oxalic) prevent long-term decline in conductivity. Xylem embolism can be quantified by expressing the initial conductivity as a percentage of the maximum obtained after flow-impeding air emboli have been removed by repeated high-pressure (175 kPa) flushes. Correlation between microbial contamination and declining conductivity suggests that long-term (> 4h) declines are caused by microbial growth within the vessels. Unpredictable trends in short-term (< 4h) measurements may be caused by movements of air emboli in vessels and/or participate matter.

Vulnerability of xylem conduits to cavitation and embolism was compared in two species of Rhizophoraceae, the mangrove Rhizophora mangle L. and the tropical moist-forest Cassipourea elliptica (Sw.) Poir. Cavitation (water column breakage preceeding embolism) was monitored by ultrasonic detection; embolism was quantified by its reduction of xylem hydraulic conductivity. Acoustic data were not predictive of loss in hydraulic conductivity, probably because signals from cavitating vessels were swamped by more numerous ones from cavitating fibers. Rhizophora mangle was the less vulnerable to embolism of the two species, losing 80% of its hydraulic conductivity between – 6.0 and – 7.0 MPa. Cassipourea elliptica lost conductivity in linear proportion to decreasing xylem pressure from – 0.5 to – 7.0 MPa. Species vulnerability correlated closely with physiological demands of habitat; the mangrove Rhizophora mangle had field xylem pressures between – 2.5 and – 4.0 MPa. whereas the minimum for Cassipourea elliptica was – 1.6 MPa. Differences in vulnerability between species could be accounted for by differences in the measured air permeability of intervessel pit membranes. According to this explanation, embolism occurs when air enters a water-filled vessel from a neighboring air-filled one via pores in shared pit membranes.

In this paper we report measurements concerning the conductivity of water and ions and the interaction between the two in excised pieces of xylem of red maple stems under various conditions. We have also demonstrated that it is possible to detect the flow of solutions through the stems of maple by measuring the degree of interaction between the flow of water and ions. In this technique we apply voltage pulses of ± V volts across a length of stem and detect the unequal current pulses resulting from the greater frictional drag when current (which is carried primarily by cations) is flowing against the water stream than when flowing with the water stream. The hydraulic conductivity of recent maple sapwood ranges from 30 to 90 cm3 sr1 cm-2 (J cm"3)"1 cm; in 2 ml KCl the electrical conductivity is roughly 3 x 10"4 mho cm-1 and the electro-kinetic cross coefficient is roughly 4x 10~5 A cm-2 (J cm"3)"1 cm.

A standing-gradient osmotic-flow model has been proposed by others to explain the apparently active water uptake in excised maize roots. An alternative model is proposed which incor porates electrogenic coupling between the fluxes of anions and cations and electro-osmotic coupling between the fluxes of cations and water in the cell walls of roots. This model does not exclude the contribution (positive or negative) of a standing-gradient osmotic flow. In recent years the ionic relations of excised roots have been somewhat extensively studied because the content of the xylem will exude from the cut end and is accessible to measurements. In an exuding root experiment one can (1) measure the rate of exudation, (2) measure the concentration and composition of the exudate, (3) vary the tonicity and composition of the bathing medium and measure the transient and steady-state effects on exudation, (4) measure the net salt uptake from the bathing medium, and (5) measure the electrical potential difference between the exudate and the bathing medium. A number of workers have measured the exudation rate, Jv (cm3 s_1 per cm2 of root surface), as a function of the difference between the osmolar concentration of the exudate, Cf, and the bathing medium, C°s (House and Findlay, 1966a; Anderson and Reilly, 1968a, b; Anderson and Collins, 1969). It has always been found that the exudation process seems to have two basic components; (1) an 'active' flux rate which is independent of water potential differences, °, and (2) a purely osmotic component oLPRT(C*—C°s) where a and LP are the reflection coefficient and hydraulic conductivity of the bulk root respectively. The equation governing exudation is then ^ = aLpBT{Cx_co)+^ (1) The values for oLP (cm3 s_1 cm-2 bar-1) and cf>° (cm3 s_1 cm-2) reported in the literature for Zea mays roots are 1-8 x 10~7 and 1-2 X 10'7 respectively (Anderson and Collins, 1969), 1*4 X 10~7 and 1-8 X 10~7 respectively (Anderson and Reilly, 1968a), and 5-7 X 10~7 and 1-3 X 10~7 respectively (House and Findlay, 1966a). Anderson, Aikman, and Meiri (1970) have recently argued that there may not be an active component of water transport in roots and they have demonstrated in theory how the entire exudation process could be osmotic. Their argument for setting = 0 is, however, ultimately circular.

We discuss the relationship between the dynamically changing tension gradients required to move water rapidly through the xylem conduits of plants and the proportion of conduits lost through embolism as a result of water tension. We consider the implications of this relationship to the water relations of trees. We have compiled quantitative data on the water relations, hydraulic architecture and vulnerability of embolism of four widely different species: Rhizophora mangle, Cassipourea elliptica, Acer saccharum, and Thuja occidentalis. Using these data, we modeled the dynamics of water flow and xylem blockage for these species. The model is specifically focused on the conditions required to generate ;runaway embolism,' whereby the blockage of xylem conduits through embolism leads to reduced hydraulic conductance causing increased tension in the remaining vessels and generating more tension in a vicious circle. The model predicted that all species operate near the point of catastrophic xylem failure due to dynamic water stress. The model supports Zimmermann's plant segmentation hypothesis. Zimmermann suggested that plants are designed hydraulically to sacrifice highly vulnerable minor branches and thus improve the water balance of remaining parts. The model results are discussed in terms of the morphology, hydraulic architecture, eco-physiology, and evolution of woody plants.

A simple steady state iterative solution of Munch pressure-flow in unbranched sieve tubes containing only water and sucrose is derived. The iterative equations can be solved on a programmable desk calculator. Solutions are presented for steady state transport with specific mass transfer rates up to 1.5 x 10(-5) mole second(-1) centimeters(-2) (= 18.5 grams hour(-1) centimeters(-2)) over distances in excess of 50 meters. The calculations clearly indicate that a Munch pressure-flow system can operate over long distances provided (a) the sieve tube is surrounded by a semipermeable membrane; (b) sugars are actively loaded in one region and unloaded at another; (c) the sieve pores are unblocked so that the sieve tube hydraulic conductivity is high (around 4 centimeters(2) second(-1) bar(-1)); (d) the sugar concentration is kept high (around one molar in the source region); and (e) the average sap velocity is kept low (around 20-50 centimeters hour(-1)). The dimensions of sieve cells in several species of plants are reviewed and sieve tube hydraulic conductivities are calculated; the values range from 0.2 to 20 centimeters(2) second(-1) bar(-1). For long distance pressure-flow to occur, the hydraulic conductivity of the sieve cell membranes must be about 5 x 10(-7) centimeters second(-1) bar(-1) or greater.