Rare pits, large vessels and extreme vulnerability to cavitation in a ring-porous tree species

? The rare pit hypothesis predicts that the extensive inter-vessel pitting in large early-wood vessels of ring-porous trees should render many of these vessels extremely vulnerable to cavitation by air-seeding. This prediction was tested in Quercus gambelii. ? Cavitation was assessed from native hydraulic conductivity at field sap tension and in dehydrated branches. Single-vessel air injections gave air-seeding pressures through vessel files; these data were used to estimate air-seeding pressures for inter-vessel walls and pits. ? Extensive cavitation occurred at xylem sap tensions below 1 MPa. Refilling occurred below 0.5 MPa and was inhibited by phloem girdling. Remaining vessels cavitated over a wide range to above 4 MPa. Similarly, 40% of injected vessel files air-seeded below 1.0 MPa, whereas the remainder seeded over a wide range exceeding 5 MPa. Inter-vessel walls averaged 1.02 MPa air-seeding pressure, similar and opposite to the mean cavitation tension of 1.22 MPa. Consistent with the rare pit hypothesis, only 7% of inter-vessel pits were estimated to air-seed by 1.22 MPa. ? The results confirm the rare pit prediction that a significant fraction of large vessels in Q. gambelii experience high probability of failure by air-seeding.     

A model of bubble growth leading to xylem conduit embolism

The dynamics of a gas bubble inside a water conduit after a cavitation event was modeled. A distinction was made between a typical angiosperm conduit with a homogeneous pit membrane and a typical gymnosperm conduit with a torus-margo pit membrane structure. For conduits with torus-margo type pits pit membrane deflection was also modeled and pit aspiration, the displacement of the pit membrane to the low pressure side of the pit chamber, was found to be possible while the emboli was still small. Concurrent with pit aspiration, the high resistance to water flow out of the conduit through the cell walls or aspirated pits will make the embolism process slow. In case of no pit aspiration and always for conduits with homogeneous pit membranes, embolism growth is more rapid but still much slower than bubble growth in bulk water under similar water tension. The time needed for the embolism to fill a whole conduit was found to be dependent on pit and cell wall conductance, conduit radius, xylem water tension, pressure rise in adjacent conduits due to water freed from the embolising conduit, and the rigidity and structure of the pits in the case of margo-torus type pit membrane. The water pressure in the conduit hosting the bubble was found to occur almost immediately after bubble induction inside a conduit, creating a sudden tension release in the conduit, which can be detected by acoustic and ultra-acoustic monitoring of xylem cavitation.     

Functional and ecological xylem anatomy

Cohesion-tension transport of water is an energetically efficient way to carry large amounts of water from the roots up to the leaves. However, the cohesion-tension mechanism places the xylem water under negative hydrostatic pressure (P_x), rendering it susceptible to cavitation. There are conflicts among the structural requirements for minimizing cavitation on the one hand vs maximizing efficiency of transport and construction on the other. Cavitation by freeze-thaw events is triggered by in situ air bubble formation and is much more likely to occur as conduit diameter increases, creating a direct conflict between conducting efficiency and sensitivity to freezing induced xylem failure. Temperate ring-porous trees and vines with wide diameter conduits tend to have a shorter growing season than conifers and diffuse-porous trees with narrow conduits. Cavitation by water stress occurs by air seeding at interconduit pit membranes. Pit membrane structure is at least partially uncoupled from conduit size, leading to a much less pronounced trade-off between conducting efficiency and cavitation by drought than by freezing. Although wider conduits are generally more susceptible to drought-induced cavitation within an organ, across organs or species this trend is very weak. Different trade-offs become apparent at the level of the pit membranes that interconnect neighbouring conduits. Increasing porosity of pit membranes should enhance conductance but also make conduits more susceptible to air seeding. Increasing the size or number of pit membranes would also enhance conductance, but may weaken the strength of the conduit wall against implosion. The need to avoid conduit collapse under negative pressure creates a significant trade-off between cavitation resistance and xylem construction cost, as revealed by relationships between conduit wall strength, wood density and cavitation pressure. Trade-offs involving cavitation resistance may explain the correlations between wood anatomy, cavitation resistance, and the physiological range of negative pressure experienced by species in their native habitats.     

Cavitation resistance of peduncle,petiole and stem is correlated with bordered pit dimensions in Magnolia grandiflora

Variation in resistance of xylem to embolism among flowers, leaves, and stems strongly influences the survival and reproduction of plants. However, little is known about the vulnerability to xylem embolism under drought stress and their relationships to the anatomical traits of pits among reproductive and vegetative organs. In this study, we investigated the variation in xylem vulnerability to embolism in peduncles, petioles, and stems in a woody plant, Magnolia grandiflora. We analyzed the relationships between water potentials that induced 50% embolism (P₅₀) in peduncles, petioles, and stems and the conduit pit traits hypothesized to influence cavitation resistance. We found that peduncles were more vulnerable to cavitation than petioles and stems, supporting the hypothesis of hydraulic vulnerability segmentation that leaves and stems are prioritized over flowers during drought stress. Moreover, P₅₀ was significantly correlated with variation in the dimensions of inter-vessel pit apertures among peduncles, petioles and stems. These findings highlight that measuring xylem vulnerability to embolism in reproductive organs is essential for understanding the effect of drought on plant reproductive success and mortality under drought stress.     

The hydraulic architecture of Juniperus communis L. ssp. communis: shrubs and trees compared

Juniperus communis ssp. communis can grow like a shrub or it can develop a tree-like habit. In this study, the hydraulic architecture of these contrasting growth forms was compared. We analysed the hydraulic efficiency (leaf-specific conductivity, k(l); specific conductivity, k(s); Huber value, HV) and the vulnerability to cavitation (the water potential corresponding to a 50% loss of conductivity, Psi(50)), as well as anatomical parameters [mean tracheid diameter, d; mean hydraulic diameter, d(h); cell wall reinforcement (t/b)(h)(2)] of shrub shoots, tree stems and tree branches. Shrub shoots were similar to tree branches (especially to lower branches) in growth form and conductivity (k(l) = 1.93 +/- 0.11 m(2) s(-1) MPa(-1) 10(-7), k(s) = 5.71 +/- 0.19 m(2) s(-1) MPa(-1) 10(-4)), but were similar to tree stems in their vulnerability to cavitation (Psi(50) = -5.81 +/- 0.08 MPa). Tree stems showed extraordinarily high k(l) and k(s) values, and HV increased from the base up. Stem xylem was more vulnerable to cavitation than branch xylem, where Psi(50) increased from lower (Psi(50) = -6.44 +/- 0.19 MPa) to upper branches (Psi(50) = -5.98 +/- 0.13 MPa). Conduit diameters were correlated with k(l) and k(s). Data indicate that differences in hydraulic architecture correspond to changes in growth form. In some aspects, the xylem hydraulics of tree-like Juniperus communis differs from that of other coniferous tree species.     

A carbon cost-gain model explains the observed patterns of xylem safety and efficiency

Efficient water transport from the soil to the leaves is essential for plant function, while building and maintaining the water transport structure in the xylem require a major proportion of the assimilated carbon of the tree. Xylem transport also faces additional challenges as water in the xylem is under tension and therefore cavitation cannot be completely avoided. We constructed a model that calculates the xylem structure that maximizes carbon-use efficiency while simultaneously taking into account pit structure in increasing the resistance to water transport and constricting the spreading of embolisms. The optimal xylem structure predicted by the model was found to correspond well to the generally observed trends: xylem conduits grew in size from the apex towards the base while simultaneously decreasing in number, and vulnerability to cavitation increased with conduit size. These trends were caused primarily by the axial water potential gradient in the xylem. The pits have to be less porous near the apex where water potential is lower to restrict the spreading of embolisms, while whole-plant carbon-use efficiency demands that conduit size decreases and conduit number increases simultaneously. The model predictions remained qualitatively the same regardless of the exact optimality criterion used for defining carbon-use efficiency.     

Cavitation fatigue. Embolism and refilling cycles can weaken the cavitation resistance of xylem

Although cavitation and refilling cycles could be common in plants, it is unknown whether these cycles weaken the cavitation resistance of xylem. Stem or petiole segments were tested for cavitation resistance before and after a controlled cavitation-refilling cycle. Cavitation was induced by centrifugation, air drying of shoots, or soil drought. Except for droughted plants, material was not significantly water stressed prior to collection. Cavitation resistance was determined from "vulnerability curves" showing the percentage loss of conductivity versus xylem pressure. Two responses were observed. "Resilient" xylem (Acer negundo and Alnus incana stems) showed no change in cavitation resistance after a cavitation-refilling cycle. In contrast, "weakened" xylem (Populus angustifolia, P. tremuloides, Helianthus annuus stems, and Aesculus hippocastanum petioles) showed considerable reduction in cavitation resistance. Weakening was observed whether cavitation was induced by centrifugation, air dehydration, or soil drought. Observations from H. annuus showed that weakening was proportional to the embolism induced by stress. Air injection experiments indicated that the weakened response was a result of an increase in the leakiness of the vascular system to air seeding. The increased air permeability in weakened xylem could result from rupture or loosening of the cellulosic mesh of interconduit pit membranes during the water stress and cavitation treatment.     

Does acclimation in cavitation resistance due to mechanical perturbation support the pit area or conduit reinforcement hypotheses in Phaseolus vulgaris?

Two Phaseolus vulgaris L. cultivars were exposed to reduced water and stem mechanical perturbation treatments (flexing) to determine if acclimation to these treatments induced hydraulic changes, altered cavitation resistance and changed stem mechanical properties. Additionally, this study sought to determine if changes in cavitation resistance would support the pit area or conduit reinforcement hypotheses. Flexing reduced biomass, leaf area, xylem vessel area and hydraulic conductivity. One cultivar had greater measures of stem strength and cavitation resistance. Flexing increased cavitation resistance (P₅₀) but did not increase Young's modulus, rigidity or flexural strength on dried stems. Stem rigidity and basal diameter were correlated with leaf mass. The ratio of conduit wall thickness to span [(t/b)_h²] increased under high water and flexing treatments while rigidity decreased for one cultivar exposed to both flexing and lower water suggesting an inability to compensate for two simultaneous stresses. Although P₅₀ was not correlated with measures of mechanical strength, P₅₀ was correlated with vessel diameter, consistent with the pit area hypothesis. This study confirmed that mechanical perturbation can impact xylem structural properties and result in altered plant water flow characteristics and cavitation resistance. Long‐term hydraulic acclimation in these herbaceous annuals was constrained by similar tradeoffs that constrain hydraulic properties across species.     

Cavitation fatigue and its reversal in sunflower (Helianthus annuus L.)

"Cavitation fatigue" is the increased susceptibility of a xylem conduit to cavitation as a result of its prior cavitation. It was investigated whether cavitation fatigue induced in vivo could be repaired in intact plants. Sunflowers (Helianthus annuus L.) were subjected to soil drought in the greenhouse. Native embolism and vulnerability to cavitation was measured in well-watered controls and after 5 d and 10 d of controlled drought. A dramatic cavitation fatigue was observed where droughted xylem that was refilled in the laboratory developed up to 60 PLC (percentage loss of hydraulic conductivity) at -1 MPa versus only 5.2 PLC in non-droughted controls. Rewatered plants showed the complete reversal of cavitation fatigue over 4 d. Reversal of fatigue was correlated with the refilling of embolized vessels in the intact plants (r(2)=0.91, P<0.01), suggesting that xylem transport to fatigued vessels was required for their repair. The in vivo reversal of fatigue was partially duplicated in excised stem segments by perfusing them with root exudates from droughted (DR) and well-watered (WW) plants. The DR exudate had a greater effect, and this was associated with a greater pH in the DR versus WW saps, but there was no difference in total cation concentration. Perfusions with 2 mM CaCl(2) and KCl solutions also partially reversed cavitation fatigue as opposed to no effect with deionized water, suggesting a role of ions in addition to a pH effect. It is suspected that fatigue is caused by stretching and partial disruption of linkages between cellulose microfibrils in inter-conduit pit membranes during air seeding, and that the reversal of fatigue involves restoring these linkages by ingredients in xylem sap.     

Xylem conduction and cavitation in Hevea brasiliensis

Clones of Hevea were studied in an attempt to discover the reasonsfor differences in the hydraulic performance of xylem. Differencesbetween clones were determined, including hydraulic conductivityand conduit width and length distributions. However, it hasproved difficult to reconcile anatomical differences with physiologicalperformance for use in future plant breeding programmes. When leaf relative water content (RWC) had been reduced fromabout 95% to 85%, the hydraulic conductivity of petioles decreasedsharply to about 40% of the initial value. This value correspondedwith xylem sap tensions of 1.8–2.0 MPa. Acoustic detectionexperiments revealed that this reduction in hydraulic conductivitycoincided with the greatest occurrence of cavitation. It seemsinescapable that the reduction in hydraulic conductivity wascaused by embolization; thereafter gas bubbles blocked the flowof water inside many of the conduits. There was some indicationthat eventually such bubbles might be dissolved, because thehydraulic conductivity increased again if specimens were fullyrehydrated. Apparently, the incidence of cavitation coincides with the entryof gas bubbles via ultramicroscopic pores into the conduitsthrough the walls according to the air-seeding hypothesis. Whena petiolate leaf is tested in a pressure chamber it is impossibleto make satisfactory measurements of a balancing pressure beyondc. 1.8–2.0 MPa, because air bubbles, mixed with sap andescaping from the conduits, form a persistent froth. Xylem transportin Hevea seems to be disrupted relatively easily under waterstress which is a feature of other tropical species adaptedto rainforest–type environments Key words: Hevea, xylem, cavitation, conduit, hydraulic conductivity     

Poplar vulnerability to xylem cavitation acclimates to drier soil conditions

Xylem vulnerability to cavitation differs between tree species according to their drought resistance, more xerophilous species being more resistant to xylem cavitation. Variability in xylem vulnerability to cavitation is also found within species, especially between in situ populations. The origin of this variability has not been clearly identified. Here we analyzed the response of xylem hydraulic traits of Populus tremula×Populus alba trees to three different soil water regimes. Stem xylem vulnerability was scored as the xylem water potential causing 12, 50 and 88% loss of conductivity (P₁₂, P₅₀ and P₈₈). Vulnerability to cavitation was found to acclimate to growing conditions under different levels of soil water content, with P₅₀ values of ?1.82, ?2.03 and ?2.45 MPa in well‐watered, moderately water‐stressed and severely water‐stressed poplars, respectively. The value of P₁₂, the xylem tension at which cavitation begins, was correlated with the lowest value of midday leaf water potential (ψm) experienced by each plant, the difference between the two parameters being approximately 0.5 MPa, consistent with the absence of any difference in embolism level between the different water treatments. These results support the hypothesis that vulnerability to cavitation is a critical trait for resistance to drought. The decrease in vulnerability to cavitation under growing conditions of soil drought was correlated with decreased vessel diameter, increased vessel wall thickness and a stronger bordered pit field (t/b)². The links between these parameters are discussed.     

Functional coordination between leaf gas exchange and vulnerability to xylem cavitation in temperate forest trees

We examined functional coordination among stem and root vulnerability to xylem cavitation, plant water transport characteristics and leaf traits in 14 co-occurring temperate tree species. Relationships were evaluated using both traditional cross-species correlations and phylogenetically independent contrast (PIC) correlations. For stems, the xylem tension at which 50% of hydraulic conductivity was lost (psi50) was positively associated (P < 0.001) with specific conductivity (K(S)) and with mean hydraulically weighted xylem conduit diameter (D(h-w)), but was only marginally (P = 0.06) associated with leaf specific conductivity (K(L)). The PIC correlation for each of these relationships, however, was not statistically significant. There was also no relationship between root psi50 and root K(S) in either cross-species or PIC analysis. Photosynthetic rate (A) and stomatal conductance (g(s)) were strongly and positively correlated with root psi50 in the cross-species analysis (P < 0.001), a relationship that was robust to phylogenetic correction (P < 0.01). A and g(s) were also positively correlated with stem psi50 in the cross-species analysis (P = 0.02 and 0.10, respectively). However, only A was associated with stem psi50 in the PIC analysis (P = 0.04). Although the relationship between vulnerability to cavitation and xylem conductivity traits within specific organs (i.e. stems and roots) was weak, the strong correlation between g(s) and root psi50 across species suggests that there is a trade-off between vulnerability to cavitation and water transport capacity at the whole-plant level. Our results were therefore consistent with the expectation of coordination between vulnerability to xylem cavitation and the regulation of stomatal conductance, and highlight the potential physiological and evolutionary significance of root hydraulic properties in controlling interspecific variation in leaf function.     

Analysis of freeze-thaw embolism in conifers. The interaction between cavitation pressure and tracheid size

Ice formation in the xylem sap produces air bubbles that under negative xylem pressures may expand and cause embolism in the xylem conduits. We used the centrifuge method to evaluate the relationship between freeze-thaw embolism and conduit diameter across a range of xylem pressures (Px) in the conifers Pinus contorta and Juniperus scopulorum. Vulnerability curves showing loss of conductivity (embolism) with Px down to -8 MPa were generated with versus without superimposing a freeze-thaw treatment. In both species, the freeze-thaw plus water-stress treatment caused more embolism than water stress alone. We estimated the critical conduit diameter (Df) above which a tracheid will embolize due to freezing and thawing and found that it decreased from 35 microm at a Px of -0.5 MPa to 6 microm at -8 MPa. Further analysis showed that the proportionality between diameter of the air bubble nucleating the cavitation and the diameter of the conduit (kL) declined with increasingly negative Px. This suggests that the bubbles causing cavitation are smaller in proportion to tracheid diameter in narrow tracheids than in wider ones. A possible reason for this is that the rate of dissolving increases with bubble pressure, which is inversely proportional to bubble diameter (La Place's law). Hence, smaller bubbles shrink faster than bigger ones. Last, we used the empirical relationship between Px and Df to model the freeze-thaw response in conifer species.     

Xylem function of arid-land shrubs from California, USA: an ecological and evolutionary analysis

Xylem traits were examined among 22 arid-land shrub species, including measures of vessel dimensions and pit area. These structural measures were compared with the xylem functional traits of transport efficiency and safety from cavitation. The influence of evolution on trait relationships was examined using phylogenetic independent contrasts (PICs). A trade-off between xylem safety and efficiency was supported by a negative correlation between vessel dimensions and cavitation resistance. Pit area was correlated with cavitation resistance when cross species data were examined, but PICs suggest that these traits have evolved independently of one another. Differences in cavitation resistance that are not explained by pit area may be related to differences in pit membrane properties or the prevalence of tracheids, the latter of which may alter pit area through the addition of vessel-to-tracheid pits or through changes in xylem conduit connectivity. Some trait relationships were robust regardless of species ecology or evolutionary history. These trait relationships are likely to be the most valuable in predictive models that seek to examine anatomical and functional trait relationships among extant and fossil woods and include the relationship among hydraulic conductivity and vessel diameter, between vessel diameter and vessel length, and between hydraulic conductivity and wood density.     

New evidence for large negative xylem pressures and their measurement by the pressure chamber method

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.     

Drought‐induced xylem pit membrane damage in aspen and balsam poplar

Drought induces an increase in a tree's vulnerability to a loss of its hydraulic conductivity in many tree species, including two common in western Canada, trembling aspen (Populus tremuloides) and balsam poplar (Populus balsamifera). Termed ‘cavitation fatigue’ or ‘air‐seeding fatigue’, the mechanism of this phenomenon is not well understood, but hypothesized to be a result of damage to xylem pit membranes. To examine the validity of this hypothesis, the effect of drought on the porosity of pit membranes in aspen and balsam poplar was investigated. Controlled drought and bench dehydration treatments were used to induce fatigue and scanning electron microscopy (SEM) was used to image pit membranes for relative porosity evaluations from air‐dried samples after ethanol dehydration. A significant increase in the diameter of the largest pore was found in the drought and dehydration treatments of aspen, while an increase in the percentage of porous pit membranes was found in the dehydration treatments of both species. Additionally, the location of the largest pore per pit membrane was observed to tend toward the periphery of the membrane.     

Testing hypotheses that link wood anatomy to cavitation resistance and hydraulic conductivity in the genus Acer

? Vulnerability to cavitation and conductive efficiency depend on xylem anatomy. We tested a large range of structure-function hypotheses, some for the first time, within a single genus to minimize phylogenetic 'noise' and maximize detection of functionally relevant variation. ? This integrative study combined in-depth anatomical observations using light, scanning and transmission electron microscopy of seven Acer taxa, and compared these observations with empirical measures of xylem hydraulics. ? Our results reveal a 2 MPa range in species' mean cavitation pressure (MCP). MCP was strongly correlated with intervessel pit structure (membrane thickness and porosity, chamber depth), weakly correlated with pit number per vessel, and not related to pit area per vessel. At the tissue level, there was a strong correlation between MCP and mechanical strength parameters, and some of the first evidence is provided for the functional significance of vessel grouping and thickenings on inner vessel walls. In addition, a strong trade-off was observed between xylem-specific conductivity and MCP. Vessel length and intervessel wall characteristics were implicated in this safety-efficiency trade-off. ? Cavitation resistance and hydraulic conductivity in Acer appear to be controlled by a very complex interaction between tissue, vessel network and pit characteristics.     

Freezing of xylem sap without cavitation

Freezing of stem sections and entire twigs of hemlock (Tsuga canadensis) has been demonstrated to occur without increasing the resistance to the movement of water through the frozen part after rewarming. This was interpreted to mean that freezing did not produce cavitation in the xylem sap even though A) the sap was unquestionably frozen; B) it contained dissolved gases; and C) it was under tension before freezing and after. Freezing stem sections of some other evergreen gymnosperms during the summer again produced no evidence for cavitation of the xylem sap. On the other hand, freezing stem sections of some angiosperms invariably increased the resistance to sap flow leading to wilting and death in a few hours when the sap tension was at normal daytime values at the time of freezing. These results were interpreted to mean that the bordered pits on the tracheids of gymnosperms function to isolate the freezing sap in each tracheid so that the expansion of water upon freezing not only eliminates any existing tension but also develops positive pressure in the sap. Dissolved gases frozen out of solution may then be redissolved under this positive pressure as melting occurs. As the bubbles are reduced in size by this ice pressure developed in an isolated tracheid, further pressure is applied by the surface tension of the water against air. If the bubbles are redissolved or are reduced to sufficient small size by the time the tension returns to the sap as the last ice crystals melt, then the internal pressure from surface tension in any existing small bubbles may exceed the hydrostatic tension of the melted sap and the bubbles cannot expand and will continue to dissolve.     

Different Aspects of Cavitation Resistance in Ceratonia siliqua, a Drought-Avoiding Mediterranean Tree

Potted plants of Ceratonia siliqua L., growing in a greenhouse,were used to detect xylem cavitation (in terms of ultrasoundacoustic emissions AE) in internodes and node-to-petiole (N-P)junctions, after different periods of drought (9, 16 and 23d). Diurnal AE were only 100 in internodes of watered (W) plantsbut 320, 1250 and 2460 in 9-, 16- and 23-d stressed ones. InN-P junctions, AE were only 15 to 20% with respect to internodes. Stem perfusion with dye allowed measurement of the percentageof xylem conduit transverse area blocked by cavitation. Thiswas 2% in internodes of W-plants and 5.2, 13.8 and 40.4% inthose of 9-, 16- and 23-d stressed ones. In N-P junctions, 18.5%of the xylem conduit transverse area was blocked in the 23-dstressed plants only. The major resistance to cavitation exhibitedby the N-P junctions is interpreted in terms of their greaternumber of narrow xylem conduits. The percentage of blocked xylemconduits within a range of diameters showed that the narrowera xylem conduit, the less likely it was that cavitation wouldoccur. After rewatering, the release of the xylem blockage causedby cavitation occurred within 2 h. Our data suggest that C.siliqua can be considered to be a cavitation avoider, especiallyin its stem-to-leaf transition zones. Ceratonia siliqua L., Carob tree, cavitation avoidance, xylem architecture, ultrasonic acoustic emissions     

Safety and efficiency conflicts in hydraulic architecture: scaling from tissues to trees

Tree hydraulic architecture exhibits patterns that propagate from tissue to tree scales. A challenge is to make sense of these patterns in terms of trade-offs and adaptations. The universal trend for conduits per area to decrease with increasing conduit diameter below the theoretical packing limit may reflect the compromise between maximizing the area for conduction versus mechanical support and storage. Variation in conduit diameter may have two complementary influences: one being compromises between efficiency and safety and the other being that conduit tapering within a tree maximizes conductance per growth investment. Area-preserving branching may be a mechanical constraint, preventing otherwise more efficient top-heavy trees. In combination, these trends beget another: trees have more, narrower conduits moving from trunks to terminal branches. This pattern: (1) increases the efficiency of tree water conduction; (2) minimizes (but does not eliminate) any hydraulic limitation on the productivity or tissue growth with tree height; and (3) is consistent with the scaling of tree conductance and sap flow with size. We find no hydraulic reason why tree height should scale with a basal diameter to the two-thirds power as recently claimed; it is probably another mechanical constraint as originally proposed. The buffering effect of capacitance on the magnitude of transpiration-induced xylem tension appears to be coupled to cavitation resistance, possibly alleviating safety versus efficiency trade-offs.