Limitation of transpiration by hydraulic conductance and xylem cavitation in Betula occidentalis

The extent to which stomatal conductance (g_s) was capable of responding to reduced hydraulic conductance (k)and preventing cavitation-inducing xylem pressures was evaluated in the small riparian tree, Betula occidentalis Hook. We decreased k by inducing xylem cavitation in shoots using an air-injection technique. From 1 to 18 d after shoot injection we measured midday transpiration rate (E), g_s, and xylem pressure (Ψ_p-xylem) on individual leaves of the crown. We then harvested the shoot and made direct measurements of k from the trunk (2–3 cm diameter) to the distal tip of the petioles of the same leaves measured for E and g_s. The k measurement was expressed per unit leaf area (k_l, leaf-specific conductance). Leaves measured within 2 d of shoot injection showed reduced g_s and E relative to non-injected controls, and both parameters were strongly correlated with k_l At this time, there was no difference in leaf Ψ_p-xylem between injected shoots and controls, and leaf Ψ_p-xylem was not significantly different from the highest cavitation-inducing pressure (Ψ_p-cav) in the branch xylem (-1.43 ± 0.029 MPa, n=8). Leaves measured 7–18 d after shoots were injected exhibited a partial return of g_s and E values to the control range. This was associated with a decrease in leaf Ψ_p-xylem below Ψ_p-cav and loss of foliage. The results suggest the stomata were incapable of long-term regulation of E below control values and that reversion to higher E caused dieback via cavitation.     

The relationship between xylem conduit diameter and cavitation caused by freezing

The centrifuge method for measuring the resistance of xylem to cavitation by water stress was modified to also account for any additional cavitation that might occur from a freeze-thaw cycle. A strong correlation was found between cavitation by freezing and mean conduit diameter. On the one extreme, a tracheid-bearing conifer and diffuse-porous angiosperms with small-diameter vessels (mean diameter <30 μm) showed no freezing-induced cavitation under modest water stress (xylem pressure = −0.5 MPa), whereas species with larger diameter vessels (mean >40 μm) were nearly completely cavitated under the same conditions. Species with intermediate mean diameters (30–40 μm) showed partial cavitation by freezing. These results are consistent with a critical diameter of 44 μm at or above which cavitation would occur by a freeze–thaw cycle at −0.5 MPa. As expected, vulnerability to cavitation by freezing was correlated with the hydraulic conductivity per stem transverse area. The results confirm and extend previous reports that small-diameter conduits are relatively resistant to cavitation by freezing. It appears that the centrifuge method, modified to include freeze–thaw cycles, may be useful in separating the interactive effects of xylem pressure and freezing on cavitation.     

Diurnal and seasonal variation in root xylem embolism in neotropical savanna woody species: impact on stomatal control of plant water status

Vulnerability to water-stress-induced embolism and variation in the degree of native embolism were measured in lateral roots of four co-occurring neotropical savanna tree species. Root embolism varied diurnally and seasonally. Late in the dry season, loss of root xylem conductivity reached 80% in the afternoon when root water potential (psi root) was about -2.6 MPa, and recovered to 25-40% loss of conductivity in the morning when psi root was about -1.0 MPa. Daily variation in psi root decreased, and root xylem vulnerability and capacitance increased with rooting depth. However, all species experienced seasonal minimum psi root close to complete hydraulic failure independent of their rooting depth or resistance to embolism. Predawn psi root was lower than psi soil when psi soil was relatively high (> -0.7 MPa) but became less negative than psi soil, later in the dry season, consistent with a transition from a disequilibrium between plant and soil psi induced by nocturnal transpiration to one induced by hydraulic redistribution of water from deeper soil layers. Shallow longitudinal root incisions external to the xylem prevented reversal of embolism overnight, suggesting that root mechanical integrity was necessary for recovery, consistent with the hypothesis that if embolism is a function of tension, refilling may be a function of internal pressure imbalances. All species shared a common relationship in which maximum daily stomatal conductance declined linearly with increasing afternoon loss of root conductivity over the course of the dry season. Daily embolism and refilling in roots is a common occurrence and thus may be an inherent component of a hydraulic signaling mechanism enabling stomata to maintain the integrity of the hydraulic pipeline in long-lived structures such as stems.     

Vulnerability to xylem cavitation and the distribution of Sonoran Desert vegetation

We studied 15 riparian and upland Sonoran desert species to evaluate how the limitation of xylem pressure (Ψ(x)) by cavitation corresponded with plant distribution along a moisture gradient. Riparian species were obligate riparian trees (Fraxinus velutina, Populus fremontii, and Salix gooddingii), native shrubs (Baccharis spp.), and an exotic shrub (Tamarix ramosissima). Upland species were evergreen (Juniperus monosperma, Larrea tridentata), drought-deciduous (Ambrosia dumosa, Encelia farinosa, Fouquieria splendens, Cercidium microphyllum), and winter-deciduous (Acacia spp., Prosopis velutina) trees and shrubs. For each species, we measured the "vulnerability curve" of stem xylem, which shows the decrease in hydraulic conductance from cavitation as a function of Ψ(x) and the Ψ(crit) representing the pressure at complete loss of transport. We also measured minimum in situ Ψ(x)(Ψ(xmin)) during the summer drought. Species in desert upland sites were uniformly less vulnerable to cavitation and exhibited lower Ψ(xmin) than riparian species. Values of Ψ(crit) were correlated with minimum Ψ(x). Safety margins (Ψ(xmin)-Ψ(crit)) tended to increase with decreasing Ψ(xmin) and were small enough that the relatively vulnerable riparian species could not have conducted water at the Ψ(x) experienced in upland habitats (-4 to -10 MPa). Maintenance of positive safety margins in riparian and upland habitats was associated with minimal to no increase in stem cavitation during the summer drought. The absence of less vulnerable species from the riparian zone may have resulted in part from a weak but significant trade-off between decreasing vulnerability to cavitation and conducting efficiency. These data suggest that cavitation vulnerability limits plant distribution by defining maximum drought tolerance across habitats and influencing competitive ability of drought tolerant species in mesic habitats.     

Loss of water transport capacity due to xylem cavitation in roots of two CAM succulents

Loss of axial hydraulic conductance as a result of xylem cavitation was examined for roots of the Crassulacean acid metabolism (CAM) succulents Agave deserti and Opuntia ficus-indica. Vulnerability to cavitation was not correlated with either root size or vessel diameter. Agave deserti had a mean cavitation pressure of -0.93 ± 0.08 MPa by both an air-injection and a centrifugal method compared to -0.70 ± 0.02 MPa by the centrifugal method for O. ficus-indica, reflecting the greater tolerance of the former species to low water potentials in its native habitat. Substantial xylem cavitation would occur at a soil water potential of -0.25 MPa, resulting in a predicted 22% loss of conductance for A. deserti and 32% for O. ficus-indica. For an extended drought of 3 mo, further cavitation could cause a 69% loss of conductance for A. deserti and 62% for O. ficus-indica. A model of axial hydraulic flow based upon the cavitation response of these species predicted that water uptake rates are far below the maximum possible, owing to the high root water potentials of these desert succulents. Despite various shoot adaptations to aridity, roots of A. deserti and O. ficus-indica are highly vulnerable to cavitation, which partially limits water uptake in a wet soil but helps reduce water loss to a drying soil.     

The importance of xylem constraints in the distribution of conifer species

Vulnerability of stem xylem to cavitation was measured in 10 species of conifers using high pressure air to induce xylem embolism. Mean values of air pressure required to induce a 50% loss in hydraulic conductivity (φ₅₀) varied enormously between species, ranging from a maximum of 14.2±0.6 MPa (corresponding to a xylem water potential of −14.2 MPa) in the semi-arid species Actinostrobus acuminatus to a minimum of 2.3±0.2 MPa in the rainforest species Dacrycarpus dacrydioides . Mean φ₅₀ was significantly correlated with the mean rainfall of the driest quarter within the distribution of each species. The value of φ₅₀ was also compared with leaf drought tolerance data for these species in order to determine whether xylem dysfunction during drought dictated drought response at the leaf level. Previous data describing the maximum depletion of internal CO₂ concentration (c_i) in the leaves of these species during artificial drought was strongly correlated with φ₅₀ suggesting a primary role of xylem in effecting leaf drought response. The possibility of a trade-off between xylem conductivity and xylem vulnerability was tested in a sub-sample of four species, but no evidence of an inverse relationship between φ₅₀ and either stem-area specific (K_a) or leaf-area specific conductivity (K₁) was found.     

Limitation of plant water use by rhizosphere and xylem conductance: results from a model

Hydraulic conductivity ( K ) in the soil and xylem declines as water potential ( Ψ ) declines. This results in a maximum rate of steady-state transpiration ( E _crit) and corresponding minimum leaf Ψ ( Ψ _crit) at which K has approached zero somewhere in the soil–leaf continuum. Exceeding these limits causes water transport to cease. A model determined whether the point of hydraulic failure (where K = 0) occurred in the rhizosphere or xylem components of the continuum. Below a threshold of root:leaf area ( A _R: A _L), the loss of rhizosphere K limited E _crit and Ψ _crit. Above the threshold, loss of xylem K from cavitation was limiting. The A _R: A _L threshold ranged from > 40 for coarse soils and/or cavitation-resistant xylem to < 0·20 in fine soils and/or cavitation-susceptible xylem. Comparison of model results with drought experiments in sunflower and water birch indicated that stomatal regulation of E reflected the species' hydraulic potential for extracting soil water, and that the more sensitive stomatal response of water birch to drought was necessary to avoid hydraulic failure. The results suggest that plants should be xylem-limited and near their A _R: A _L threshold. Corollary predictions are (1) within a soil type the A _R: A _L should increase with increasing cavitation resistance and drought tolerance, and (2) across soil types from fine to coarse the A _R: A _L should increase and maximum cavitation resistance should decrease.     

Capacitive effect of cavitation in xylem conduits: results from a dynamic model

Embolisms decrease plant hydraulic conductance and therefore reduce the ability of the xylem to transport water to leaves provided that embolized conduits are not refilled. However, as a xylem conduit is filled with gas during cavitation, water is freed to the transpiration stream and this transiently increases xylem water potential. This capacitive effect of embolism formation on plant function has not been explicitly quantified in the past. A dynamic model is presented that models xylem water potential, xylem sap flow and cavitation, taking into account both the decreasing hydraulic conductance and the water release effect of xylem embolism. The significance of the capacitive effect increases in relation to the decreasing hydraulic conductance effect when transpiration rate is low in relation to the total amount of water in xylem conduits. This ratio is typically large in large trees and during drought.     

Evaluation of centrifugal methods for measuring xylem cavitation in conifers, diffuse- and ring-porous angiosperms

A centrifugal method is used to measure 'vulnerability curves' which show the loss of hydraulic conductivity in xylem by cavitation. Until recently, conductivity was measured between bouts of centrifugation using a gravity-induced head. Now, conductivity can be measured during centrifugation. This 'spin' method is faster than the 'gravity' technique, but correspondence between the two has not been evaluated. The two methods were compared on the same stem segments for two conifer, four diffuse-porous, and four ring-porous species. Only 17 of 60 conductivity measurements differed, with differences in the order of 10%. When different, the spin method gave higher conductivities at the beginning of the curve and lower at the end. Pressure at 50% loss of conductivity, and mean cavitation pressure, were the same in 14 of 20 comparisons. When different, the spin method averaged 0.32 MPa less negative. Ring-porous species showed a precipitous initial drop in conductivity by both techniques. This striking pattern was confirmed by the air-injection method and native embolism measurements. Close correspondence inspires confidence in both methods, each of which has unique advantages. The observation that ring-porous species operate at only a fraction of their potential conductivity at midday demands further study.     

Stomatal control of xylem embolism

Abstract. The potential role of stomatal closure in the control of xylem embolism is investigated by means of a simple model of hydraulic flow in plants. Maintenance of a maximally efficient conducting system requires the stomata to close in an appropriate fashion as evaporative demand increases in order to prevent shoot water potentials falling below the threshold value at which cavitations occur. The model showed that the optimal stomatal behaviour required depends on soil water availability. Further analysis of the model demonstrated that there could be certain circumstances where loss of a proportion of the conducting tissue by embolisms can, perhaps surprisingly, be beneficial in terms of maximizing stomatal aperture and hence short-term productivity. The results are discussed in relation to the signals controlling stomatal aperture, and it is shown that (1) optimal control cannot be obtained using information on leaf water potential alone, and (2) information relating to soil water potential is a necessary requirement for optimal control.     

Drought resistance of Quercus pubescens as a function of root hydraulic conductance, xylem embolism and hydraulic architecture

Water relations, xylem embolism, root and shoot hydraulic conductance of both young plants in the field and potted seedlings of Quercus pubescens have been studied with the aim of investigating whether these variables may account for the well known adaptation of this oak species to arid habitats. Our data revealed that Q. pubescens is able to maintain high leaf relative water contents under water stress conditions. In fact, relative water contents measured in summer (July) did not differ from those recorded in April. This was apparently achieved by compensating water loss by an equal amount of water uptake. Such a drought avoidance strategy was made possible by the recorded high hydraulic efficiency of stems and roots under water stress. In fact, root hydraulic conductance of field-grown plants was maintained high in summer when the percentage loss of hydraulic conductance of stems was lowest. The hydraulic architecture of young plants of Q. pubescens measured in terms of partitioning of hydraulic resistances along the water pathway revealed that the highest hydraulic resistance was located in stems of the current year's growth. This hydraulic architecture is interpreted as consistent with the adaptation of Q. pubescens to arid habitats as a consequence of the recorded seasonal changes in water relation parameters as well as in root and stem hydraulics.     

Neither xylem collapse,cavitation, or changing leaf conductance drive stomatal closure in wheat

Identifying the drivers of stomatal closure and leaf damage during stress in grasses is a critical prerequisite for understanding crop resilience. Here, we investigated whether changes in stomatal conductance (g_s) during dehydration were associated with changes in leaf hydraulic conductance (K_leaf), xylem cavitation, xylem collapse, and leaf cell turgor in wheat (Triticum aestivum). During soil dehydration, the decline of g_s was concomitant with declining K_leaf under mild water stress. This early decline of leaf hydraulic conductance was not driven by cavitation, as the first cavitation events in leaf and stem were detected well after K_leaf had declined. Xylem vessel deformation could only account for <5% of the observed decline in leaf hydraulic conductance during dehydration. Thus, we concluded that changes in the hydraulic conductance of tissues outside the xylem were responsible for the majority of K_leaf decline during leaf dehydration in wheat. However, the contribution of leaf resistance to whole plant resistance was less than other tissues (<35% of whole plant resistance), and this proportion remained constant as plants dehydrated, indicating that K_leaf decline during water stress was not a major driver of stomatal closure.     

Xylem cavitation and hydraulic control of stomatal conductance in Laurel (Laurus nobilis L.)

The possible link between stomatal conductance (g_L), leaf water potential ( Ψ _L) and xylem cavitation was studied in leaves and shoots of detached branches as well as of whole plants of Laurus nobilis L. (Laurel). Shoot cavitation induced complete stomatal closure in air‐dehydrated detached branches in less than 10 min. By contrast, a fine regulation of g_L in whole plants was the consequence of Ψ _L reaching the cavitation threshold ( Ψ _CAV) for shoots. A pulse of xylem cavitation in the shoots was paralleled by a decrease in g_L of about 50%, while Ψ _L stabilized at values preventing further xylem cavitation. In these experiments, no root signals were likely to be sent to the leaves from the roots in response to soil dryness because branches were either detached or whole plants were growing in constantly wet soil. The stomatal response to increasing evaporative demand appeared therefore to be the result of hydraulic signals generated during shoot cavitation. A negative feedback link is proposed between g_L and Ψ _CAV rather than with Ψ _L itself.     

A method for measuring hydraulic conductivity and embolism in xylem

Abstract 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.     

Inter-vessel pitting and cavitation in woody Rosaceae and other vesselled plants: a basis for a safety versus efficiency trade-off in xylem transport

The hypothesis that greater safety from cavitation by air-seeding through inter-vessel pits comes at the cost of less porous pit membranes with greater flow resistance was tested . Sixteen vessel-bearing species were compared: 11 from the Rosaceae, four from other angiosperm families, and one fern. Unexpectedly, there was no relationship between pit resistance (and hence the prevailing membrane porosity) and cavitation pressure. There was, however, an inverse relationship between pit area per vessel and vulnerability to cavitation (r² = 0.75). This suggests that cavitation is caused by the rare largest membrane pore per vessel, the average size of which increases with total pit area per vessel. If safety from cavitation constrains pit membrane surface area, it also limits vessel surface area and the minimum vessel resistivity. This trade-off was consistent with an approximately three-fold increase in vessel resistivity with cavitation pressure dropping from −0.8 to −6.6 MPa. The trade-off was compensated for by a reduction in the percentage of vessel wall pitted: from 10–16% in vulnerable species to 2–4% in resistant species. Across species, end-wall pitting accounted for 53 ± 3% of the total xylem resistivity. This corresponded to vessels achieving on average 94 ± 2% of their maximum possible conductivity if vessel surface area is constrained.     

Effects of the hydraulic coupling between xylem and phloem on diurnal phloem diameter variation

Measurements of diurnal diameter variations of the xylem and phloem are a promising tool for studying plant hydraulics and xylem-phloem interactions in field conditions. However, both the theoretical framework and the experimental verification needed to interpret phloem diameter data are incomplete. In this study, we analytically evaluate the effects of changing the radial conductance between the xylem and the phloem on phloem diameter variations and test the theory using simple manipulation experiments. Our results show that phloem diameter variations are mainly caused by changes in the radial flow rate of water between the xylem and the phloem. Reducing the hydraulic conductance between these tissues decreases the amplitude of phloem diameter variation and increases the time lag between xylem and phloem diameter variation in a predictable manner. Variation in the amplitude and timing of diameter variations that cannot be explained by changes in the hydraulic conductance, could be related to changes in the osmotic concentration in the phloem.     

水分和盐分胁迫对番茄植株生长和木质部水力特性的影响

木质部是植株体内水分传输的主要通路,其水力特性的变化会影响植株的水分关系和果实的水分积累。目前关于番茄植株木质部解剖结构和水力特性对水分和盐分胁迫的响应及其与植株生长和果实含水量之间的关系尚不明确。本研究通过日光温室番茄盆栽试验,设置3个处理:对照,土壤含水量(θ)为75%～95%田间持水量(FC),初始电导率(EC)为0.398 dS·m^-1;水分胁迫,开花前θ为75%～95% FC,开花后至成熟期θ为45%～65% FC,EC为0.398 dS·m^-1;盐分胁迫,θ为75%～95% FC,EC为1.680 dS·m^-1,研究了樱桃型番茄(红宝石)和中果型番茄(北番501)植株在水分和盐分胁迫下的植株生长、果实含水量以及木质部水力特性的变化。结果表明: 与对照相比,水分和盐分胁迫下茎秆横截面积和木质部导管直径分别减小了22.0%～40.7%和10.0%～18.3%,茎秆比导水率和桁架柄比导水率分别降低了8.8%～41.1%和12.9%～28.4%,抑制了植株生长,减少了地上部鲜重、果实大小、果实鲜重和含水量,且与樱桃型番茄相比,中果型番茄的降幅更大。此外,果实含水量分别与茎秆和桁架柄比导水率呈显著正相关。综上,番茄植株在水分和盐分胁迫下木质部水力特性指标减小,生长被抑制,果实鲜重显著降低,最终导致产量降低。其中,中果型番茄相较于樱桃型番茄对水分和盐分胁迫更敏感。