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.     

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.     

Freezing-induced xylem cavitation and the northern limit of Larrea tridentata

We investigated the occurrence of freezing-induced cavitation in the evergreen desert shrub Larrea tridentata and compared it to co-occurring, winter-deciduous Prosopis velutina. Field measurements indicated that xylem sap in L. tridentata froze at temperatures below c. –5°C, and that this caused no measurable cavitation for minimum temperatures above –7°C. During the same period P. velutina cavitated almost completely. In the laboratory, we cooled stems of L. tridentata to temperatures ranging from –5 to –20°C, held them at temperature for 1 or 12 h, thawed the stems at a constant rate and measured cavitation by the decrease in hydraulic conductivity of stem segments. As observed in the field, freezing exotherms occurred at temperatures between –6.5 and –9°C and as long as temperatures were held above –11°C there was no change in hydraulic conductivity after thawing. However, when stems were cooled to between –11°C and –20°C, stem hydraulic conductivity decreased linearly with minimum temperature. Minimum temperatures between –16 and –20°C were sufficient to completely eliminate hydraulic conductance. Record (>20 year) minimum isotherms in this same range of temperatures corresponded closely with the northern limit of L. tridentata in the Mojave and Sonoran deserts.     

胡杨木质部水分传导对盐胁迫的响应与适应北大核心CSCD

解析植物木质部导水率对逆境的响应和适应对促进植物抗逆性机理研究和受损植被恢复具有重要意义。该文以荒漠河岸林建群种胡杨(Populus euphratica)为研究对象,系统分析了胡杨幼株根、茎、叶水分传输通道对不同浓度盐胁迫的响应和适应。结果表明:(1)胡杨幼株根系对盐胁迫的敏感性高于茎和叶,盐胁迫下根系生长和根尖数显著受到抑制,根木质部易于发生栓塞,导水率明显降低。(2)胡杨幼株茎木质部导水率对盐胁迫的响应依盐浓度而定,轻度(0.05 mol·L–1 Na Cl)和中度(0.15 mol·L–1 Na Cl)盐胁迫下,胡杨可以通过协调导管输水的有效性和安全性来调节木质部的导水率,维持植物正常生长;重度(0.30 mol·L–1 Na Cl)盐胁迫下,胡杨茎木质部导管输水有效性和安全性均明显降低,木质部导水率显著下降,并伴随叶片气孔导度的显著降低,从而严重抑制了胡杨的光合和生长。     

胡杨木质部水分传导对盐胁迫的响应与适应

解析植物木质部导水率对逆境的响应和适应对促进植物抗逆性机理研究和受损植被恢复具有重要意义。该文以荒漠河岸林建群种胡杨(Populus euphratica)为研究对象, 系统分析了胡杨幼株根、茎、叶水分传输通道对不同浓度盐胁迫的响应和适应。结果表明: (1)胡杨幼株根系对盐胁迫的敏感性高于茎和叶, 盐胁迫下根系生长和根尖数显著受到抑制, 根木质部易于发生栓塞, 导水率明显降低。(2)胡杨幼株茎木质部导水率对盐胁迫的响应依盐浓度而定, 轻度(0.05 mol·L^-1 NaCl)和中度(0.15 mol·L^-1 NaCl)盐胁迫下, 胡杨可以通过协调导管输水的有效性和安全性来调节木质部的导水率, 维持植物正常生长; 重度(0.30 mol·L^-1 NaCl)盐胁迫下, 胡杨茎木质部导管输水有效性和安全性均明显降低, 木质部导水率显著下降, 并伴随叶片气孔导度的显著降低, 从而严重抑制了胡杨的光合和生长。     

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.     

Intra- and inter-plant variation in xylem cavitation in Betula occidentalis

A modified version of a method that uses positive air pressures to determine the complete cavitation response of a single axis is presented. Application of the method to Betula occidentalis Hook, gave a cavitation response indistinguishable from that obtained by dehydration, thus verifying the technique and providing additional evidence that cavitation under tension occurs by air entry through interconduit pits. Incidentally, this also verified pressure-bomb estimates of xylem tension and confirmed the existence of large (i.e. >0·4 MPa) tensions in xylem, which have been questioned in recent pressure-probe studies. The air injection method was used to investigate variation within and amongst individuals of B. occidentalis. Within an individual, the average cavitation tension increased from 0·66±0·27 MPa in roots (3·9 to 10·7 mm diameter), to 1·17±0·10 MPa in trunks (12 to 16 mm diameter), to 1·36±0·04 MPa in twigs (3·9 to 5 mm diameter). Cavitation tension was negatively correlated with the hydraulically weighted mean of the vessel diameter, and was negatively correlated with the conductance of the xylem per xylem area. Native cavitation was within the range predicted from the measured cavitation response and in situ maximum xylem tensions: roots were significantly cavitated compared with minimal cavitation in trunks and twigs. Leaf turgor pressure declined to zero at the xylem tensions predicted to initiate cavitation in petiole xylem (1·5 MPa). Amongst individuals within B. occidentalis, average cavitation tension in the main axis varied from 0·90 to 1·90 MPa and showed no correlation with vessel diameter. The main axes of juveniles (2–3 years old) had significantly narrower vessel diameters than those of adults, but there was no difference in the average cavitation tension. However, juvenile xylem retained hydraulic conductance to a much higher xylem tension (3·25 MPa) than did adult xylem (2·25 MPa), which could facilitate drought survival during establishment.     

Photosynthesis in cold acclimated leaves of plants with various degrees of freezing tolerance

The objective of this study was to compare the photosynthetic changes during cold acclimation in various plant types able to acquire different degrees of freezing tolerance. Four herbaceous and six woody plants were hardened under natural or artificial conditions and – after determination of their frost resistance (LT₅₀) – the net photosynthetic rate at an ambient CO₂ of 33 Pa (P_n33), the dependencies of P_n to light and to CO₂ and the room temperature chlorophyll a fluorescence were recorded under optimal conditions. Herbaceous plants acquired freezing tolerances to temperatures between ?10 and ?15°C when hardened at temperatures around 0°C. Most leaves fully developed prior to frost hardening exhibited typical symptoms of senescence after frost hardening. In non-senescing leaves P_n33 was reduced by 15 to 50% mainly due to a reduced stomatal conductance. After hardening at temperatures around ?10°C Brassica survived down to ?24°C, but P_n33 was almost abolished as a result of disturbances in the chloroplasts. After transferring the plants to 20/15°C P_n33 recovered completely within a few days. Woody plants hardened at temperatures around 0°C tolerated – 15 to ?36°C: P_n33 was reduced by 25 to 60% and hardly recovered at 20/15°C. Hardening at ?10°C induced a tolerance of ?32 to n33 was almost totally blocked, but at 20/15°C it returned to the values of the plants hardened at 0°C within a few days. In woody plants disturbances were invariably localized in the chloroplasts. Thus, conifers, and especially Pinus cembra, can survive much lower temperatures than herbaceous plants and, at the same level of freezing tolerance, exhibit appreciably less restriction in relative P_n33.     

Scaling of xylem vessels and veins within the leaves of oak species

General models of plant vascular architecture, based on scaling of pipe diameters to remove the length dependence of hydraulic resistance within the xylem, have attracted strong interest. However, these models have neglected to consider the leaf, an important hydraulic component; they assume all leaves to have similar hydraulic properties, including similar pipe diameters in the petiole. We examine the scaling of the leaf xylem in 10 temperate oak species, an important hydraulic component. The mean hydraulic diameter of petiole xylem vessels varied by 30% among the 10 oak species. Conduit diameters narrowed from the petiole to the midrib to the secondary veins, consistent with resistance minimization, but the power function scaling exponent differed from that predicted for stems. Leaf size was an organizing trait within and across species. These findings indicate that leaf vasculature needs to be included in whole-plant scaling models, for these to accurately reflect and predict whole-plant transport and its implications for performance and ecology.     

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.     

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.     

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.     

Phosphorus nutritional effects on root hydraulic conductance,xylem water flow and flux of magnesium and calcium in squash plants

Previous studies have found that P nutrition of plants is an important factor in the uptake and translocation of Mg and Ca, and increasing root osmotic hydraulic conductance (L_o) and osmotically driven xylem exudate flow (J_v). Experiments were designed to determine if the observed changes in Mg and Ca uptake and translocation, J_v, and L_o from altered P nutrition are related or are separate functions. When six-week old squash (Cucurbita pepo L.) plants grown in perlite were treated with P levels ranging from 50 to 400 μM P for seven days, J_v and L_o increased as P treatment level increased. Xylem exudate concentrations of Mg and Ca were maintained as J_v increased, resulting in an increase in total flux of these mineral elements. The increase in Mg and Ca flux in the xylem exudate correlated with increased shoot Mg and Ca levels as P nutritional level was raised. Further studies with greenhouse grown plants indicated that the increases in J_v, L_o, and Mg and Ca flux were more responsive to changes in P nutritional level than to similar changes in levels of other anions. In hydroponically grown squash plants, xylem exudate was collected for a 20 min period after 0, 2 and 4 h in treatments of 50 and 500 μM P or after P treatment was increased from 50 to 500 μM. Immediately after nutrient solution P was increased (time 0), there was a 33% increase in J_v and a 22% increase in L_o when compared to the 50 μM P treatment. The J_v and L_o of the 50–500 μM P treatment did not equal levels of the continuous 500 μM control at time 0, but were similar after 2 and 4 h. Flux of Mg and Ca did not increase as rapidly as J_v in the 50–500 treatment indicating that regulation of Mg and Ca uptake and xylem loading by P may lag behind that of water movement. This revised version was published online in June 2006 with corrections to the Cover Date.     

Hydraulic conductance and water potential gradients in squash leaves showing mycorrhiza-induced increases in stomatal conductance

Stomatal conductance (g _s) and transpiration rates vary widely across plant species. Leaf hydraulic conductance (k _leaf) tends to change with g _s, to maintain hydraulic homeostasis and prevent wide and potentially harmful fluctuations in transpiration-induced water potential gradients across the leaf (ΔΨ _leaf). Because arbuscular mycorrhizal (AM) symbiosis often increases g _s in the plant host, we tested whether the symbiosis affects leaf hydraulic homeostasis. Specifically, we tested whether k _leaf changes with g _s to maintain ΔΨ _leaf or whether ΔΨ _leaf differs when g _s differs in AM and non-AM plants. Colonization of squash plants with Glomus intraradices resulted in increased g _s relative to non-AM controls, by an average of 27% under amply watered, unstressed conditions. Stomatal conductance was similar in AM and non-AM plants with exposure to NaCl stress. Across all AM and NaCl treatments, k _leaf did change in synchrony with g _s (positive correlation of g _s and k _leaf), corroborating leaf tendency toward hydraulic homeostasis under varying rates of transpirational water loss. However, k _leaf did not increase in AM plants to compensate for the higher g _s of unstressed AM plants relative to non-AM plants. Consequently, ΔΨ _leaf did tend to be higher in AM leaves. A trend toward slightly higher ΔΨ _leaf has been observed recently in more highly evolved plant taxa having higher productivity. Higher ΔΨ _leaf in leaves of mycorrhizal plants would therefore be consistent with the higher rates of gas exchange that often accompany mycorrhizal symbiosis and that are presumed to be necessary to supply the carbon needs of the fungal symbiont.     

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.     

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.