Role of Pressure in Xylem Transport of Coconut and Other Palms

The significance of root pressure in the transport of xylem sap has been investigated in Cocos nucifera L. and a few other palms. Despite the fact that excised palm roots can generate considerable pressures in situ, the quantity of water transported is only a small fraction of the demand resulting from transpiration. Most water transport is induced by negative pressure gradients, as in other higher plants. The development of considerable negative pressures has been demonstrated both directly and indirectly. Acoustic detection was used for the first time to monitor cavitation in water-stressed Cocos leaves. Its detection implies the ready disruption of xylem sap under these tensions. We suggest that root pressure might serve to refill cavitated xylem conduits when water is abundantly available and transpiration practically zero. However, little or no positive pressure could be demonstrated in intact palms subjected to low water stress: experimentally.     

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.     

Drought-induced increase in xylem malate and mannitol concentrations and closure of Fraxinus excelsior L. stomata

Changes in the malate and mannitol composition of ash leaf (Fraxinus excelsior L.) xylem sap were studied in response to water deficit. Xylem sap was collected by the pressure method from the petiole of leaves sampled on irrigated and non-irrigated ash seedlings. As the leaf water potential decreased from -0.3 to -3.0 MPa, there was a significant increase in malate and mannitol xylem concentrations, and a concomitant decrease in maximal stomatal conductance. The functional significance of the increased malate and mannitol concentrations was investigated by using a transpiratory bioassay with mature detached leaves which exhibited, for stomatal conductance, the typical pattern showed by expanded leaves during dark/light transitions. Supplying detached leaves with mannitol in a range of concentrations found in the xylem sap had no effect on stomatal movements, but malate, for concentrations between 0.5 and 3 mM, was effective in preventing stomatal opening. The ability of malate to inhibit stomatal opening appeared to be rather non-specific. Two structural malate analogues, citrate and aspartate or an unrelated anion, shikimate, also inhibited this process. Given the drought-induced increase in xylem malate concentrations, and the fact that the range of malate levels required to close stomata was very similar to that of the concentrations found in the xylem sap, it has been hypothesized that malate is involved in the stomatal closure of ash leaves under drying conditions.Key words: Fraxinus excelsior: L., malate, mannitol, xylem sap, stomata, water deficit.      

Sap Ascent in Vascular Plants: Challengers to the Cohesion Theory Ignore the Significance of Immature Xylem and the Recycling of Munch Water

In recent years the cohesion theory has been attacked on thegrounds that direct measurements made with the pressure probeindicate that sap tensions are much less (maximum tension approx.0.7 MPa) than indicated by parallel measurements made with themore conventional methods: osmotic methods, pressure bomb, orpsychrometer. It has also been claimed that other direct methodsdo not support the cohesion theory. Thus a re-examination usingthe Renner technique indicated sap tensions of approx. 2.5 MPa.Also an independent method based on mercury penetrometry providesevidence that sap tensions of at least 2.0 MPa can be demonstrateddirectly implying, that serious limitations arise from the pressureprobe method itself. Without tensions exceeding 2.0 MPa mangroveswould be unable to extract fresh water for transpiration fromseawater. It is suggested that the pressure probe is susceptibleto bias because it investigates the least mature xylem conduitswhile they are still under varying degrees of turgor pressureand only partially interconnected with the main xylem system.This supposition is supported by claims that the xylem sap sampledby the probe contains significant concentrations of solutes.Additionally water, supplied by reverse osmosis from the sievetubes (‘Münch water’), is continually beingliberated in the vicinity of the outermost xylem vessels hydratingthem to an atypical degree which can explain several of thediscrepancies claimed. These results, which are supported bythe work of others, demonstrate that the challenges to the cohesiontheory for the ascent of sap are ill-founded. The release ofwater from the phloem can explain not only some discrepanciesclaimed by the cohesion challengers, but also explain the refillingof cavitated xylem conduits: a hitherto unsuspected role forthe phloem transport system. Cohesion theory; sap ascent; cavitation; pressure probe; xylem transport; vessel development; recycled water; reverse osmosis     

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     

Ultrasound acoustic emissions from dehydrating leaves of deciduous and evergreen trees

While drying, detached leaves produced ultrasound acoustic emissions (UAE) comparable to emissions from stem and twig wood. Experiments on Ilex aquifolium L. showed that the main source of these signals was cavitation in the veins, to which conduits and fibres probably both contributed. Regions of the leaf blade with abundant mesophyll and only small veins emitted few signals. More signals were counted on the adaxial side of the midrib than on the abaxial one and on the proximal third than on the distal one, in accordance with the anatomical structure. Sound attenuation was pronounced. Eight species were compared with respect to cavitation behaviour, field water relations and pressure–volume curves, and the data showed differences in cumulative number of events and resistance of leaves to cavitation. Data were generally in good agreement with anatomical structure and habitat preferences. The number of signals per conduit counted on cross-sections was in some leaves much higher than unity, which suggests short xylem elements or an acoustic activity of cells other than conduits. There was no correlation between cavitation threshold or cumulative number of signals and the degree of sclerophylly; unexpectedly, there was a correlation between the cumulative number of signals at a water potential of -1.3 MPa and the bulk modulus of elasticity.     

Water storage in the wood and xylem cavitation in 1-year-old twigs of Populus deltoides Bartr.

The possible role of water expelled from cavitated xylem conduits in the rehydration of water-stressed leaves has been studied in one-year-old twigs of populus deltoides Bartr. Twigs were dehydrated in air. At desired values of leaf water potential (Ψ_l) (between near full turgor and -1.62 MPa), twigs were placed in black plastic bags for 1–2h. Leaf water content was measured every 3–5 min before bagging and every 10 min in the dark. Hydraulic conductivity and xylem cavitation were measured both in the open and in the dark. Cavitation was monitored as ultrasound acoustic emissions (AE). A critical Ψ_l value of -0.96 MPa was found, at which AE increased significantly while the leaf water deficit decreased by gain of water. Since the twigs were no longer attached to roots, it was concluded that water expelled from cavitated xylem conduits was transported to the leaves, thus contributing to their rehydration. Xylem cavitation is discussed in terms of a ‘leaf water deficit buffer mechanism’, under not very severe water stress conditions.     

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.     

Linking xylem diameter variations with sap flow measurements

Measurements of variation in the diameter of tree stems provide a rapid response, high resolution tool for detecting changes in water tension inside the xylem. Water movement inside the xylem is caused by changes in the water tension and theoretically, the sap flow rate should be directly proportional to the water tension gradient and, therefore, also linearly linked to the xylem diameter variations. The coefficient of proportionality describes the water conductivity and elasticity of the conducting tissue. Xylem diameter variation measurements could thus provide an alternative approach for estimating sap flow rates, but currently we lack means for calibration. On the other hand, xylem diameter variation measurements could also be used as a tool for studying xylem structure and function. If we knew both the water tension in the xylem and the sap flow rate, xylem conductivity and/or elasticity could be calculated from the slope of their relationship. In this study we measured diurnal xylem diameter variation simultaneously with sap flow rates (Granier-type thermal method) in six deciduous species (Acer rubrum L., Alnus glutinosa Miller, Betula lenta L., Fagus Sylvatica L. Quercus rubra L., and Tilia vulgaris L.) for 7–91 day periods during summers 2003, 2005 and 2006 and analyzed the relationship between these two measurements. We found that in all species xylem diameter variations and sap flow rate were linearly related in daily scale (daily average R ² = 0.61–0.87) but there was a significant variation in the daily slopes of the linear regressions. The largest variance in the slopes, however, was found between species, which is encouraging for finding a species specific calibration method for measuring sap flow rates using xylem diameter variations. At a daily timescale, xylem diameter variation and sap flow rate were related to each other via a hysteresis loop. The slopes during the morning and afternoon did not differ statistically significantly from each other, indicating no overall change in the conductivity. Because of the variance in the daily slopes, we tested three different data averaging methods to obtain calibration coefficients. The performance of the averaging methods depended on the source of variance in the data set and none of them performed best for all species. The best estimates of instantaneous sap flow rates were also given by different averaging methods than the best estimates of total daily water use. Using the linear relationship of sap flow rate and xylem diameter variations we calculated the conductance and specific conductivity of the soil–xylem–atmosphere water pathway. The conductance were of the order of magnitude 10⁻⁵ kg s⁻¹ MPa⁻¹ for all species, which compares well with measured water fluxes from broadleaved forests. Interestingly, because of the large sap wood area the conductance of Betula was approximately 10 times larger than in other species.     

Water stress induced cavitation and embolism in some woody plants

A comparison was made of the relative vulnerability of xylem conduits to cavitation and embolism in three species [ Thuja occidentalis L., Tsuga canadensis (L.) Carr. and Acer saccharum Marsh.]. Waterlogged samples of wood were air dehydrated while measuring relative water loss, loss of hydraulic conductance, cumulative acoustic emissions (= cavitations) and xylem water potential. Most cavitation events and loss of hydraulic conductance occurred while water potential declined from – 1 to –6 MPa. There were differences in vulnerability between species. Other people have hypothesized that large xylem conduits (e.g. vessels) should be more vulnerable to cavitations than small conduits (e.g. tracheids). Our findings are contrary to this hypothesis. Under water stress, the vessel bearing wood retained water better than tracheid bearing wood. However, within a species large conduits were more prone to cavitation than small conduits.     

High Molecular Weight Organic Compounds in the Xylem Sap of Mangroves: Implications for Long-Distance Water Transport

The rise of sap in mangroves has puzzled plant physiologists for many decades. The current consensus is that negative pressures in the xylem exist which are sufficiently high to exceed the osmotic pressure of seawater (2.5 MPa). This implies that the radial reflection coefficients of the mangrove roots are equal to unity. However, direct pressure probe measurements in xylem vessels of the roots and stems of mangrove (Rhizophora mangle) grown in the laboratory or in the field yielded below-atmospheric, positive (absolute) pressure values. Slightly negative pressure values were recorded only occasionally. Xylem pressure did not change significantly when the plants were transferred from tap water to solutions containing up to 1700 mOsmol kg^?1 NaCl. This indicates that the radial reflection coefficient of the roots for salt, and therefore the effective osmotic pressure of the external solution, was essentially zero as already reported for other halophytes. The low values of xylem tension measured with the xylem pressure probe were consistent with previously published data obtained using the vacuum/leafy twig technique. Values of xylem tension determined with these two methods were nearly two orders of magnitude smaller than those estimated for mangrove using the pressure chamber technique (?3 to ?6MPa). Xylem pressure probe measurements and staining experiments with alcian blue and other dyes gave strong evidence that the xylem vessels contained viscous, mucilage- and/or protein-related compounds. Production of these compounds resulting from wound or other artifactual reactions was excluded. The very low sap flow rates of about 20–50 cm h^?1 measured in these mangrove plants were consistent with the presence of high molecular weight polymeric substances in the xylem sap. The presence of viscous substances in the xylem sap of mangroves has the following implications for traditional xylem pressure measurement techniques, development of xylem tension, and longdistance water transport: (1) high external balancing pressures in the pressure chamber are needed to force xylem sap to the cut surface of the twig; (2) stable tensions much larger than 0.1 MPa can be developed only occasionally because viscous solutions provide nucleation sites for gas bubble formation; (3) the frequent presence of small gas bubbles in viscous solutions allows water transport by interfacial, gravity-independent streaming at gas/water interfaces and (4) the increased density of viscous solutions creates (gravity-dependent) convectional flows. Density-driven convectional flows and interfacial streaming, but also the very low radial reflection coefficient of the roots to NaCl are apparently the means by which R. mangle maintains water transport to its leaves despite the high salinity of the environment.     

What causes the differences in cavitation resistance of two shrubs? Wood anatomical explanations and reliability testing of vulnerability curves

Relationships between xylem anatomical traits and cavitation resistance have always been a major content of plant hydraulics. To know how plants cope with drought, it is extremely important to acquire detailed knowledge about xylem anatomical traits and assess the cavitation resistance accurately. This study aims to increase our knowledge in the methods determining cavitation resistance and xylem anatomical traits. We selected a semi-ring-porous species, Hippophae rhamnoides L., and a diffuse-porous species, Corylus heterophylla F., to clarify the reasons for the difference in cavitation resistance based on detailed xylem anatomical traits and reliable vulnerability curves (VCs). Both Cavitron and bench dehydration (BD) were used to construct VCs. Xylem anatomical traits, including pit membrane ultrastructure of these two species, were determined. The VCs obtained by the two different techniques were of different types for H. rhamnoides, its Cavitron VCs might be unreliable because of open-vessel artifacts. On the basis of BD VCs, H. rhamnoides showed higher cavitation resistance than C. heterophylla, and this is attributed to its low vessel connectivity as well as non-porous and thicker pit membranes.     

Seasonal ultrasonic acoustic emissions of <Emphasis Type="Italic">Quercus ilex</Emphasis> L. trees in a Mediterranean forest

Ultrasonic acoustic emissions were measured in Quercus ilex trees of a Mediterranean forest in Catalonia (NE Spain) each season from summer of 2004 to autumn of 2005. Acoustic emissions were maximum during hot and dry summer periods. Acoustic emissions started below 17% soil moisture, 0.85 RWC, and 2.5 MPa leaf water potential. They were negatively correlated with soil moisture and leaf water potential. The relationship between acoustic emissions and leaf water potential was the strongest, indicating that xylem tension is the most important factor inducing both cavitation (acoustic emissions) and a decrease in leaf water potential. Future increase of xylem cavitation derived from climate change may result in growth and survival limitations for this species in the drier southern limits of its current distribution.     

A New Theory for the Ascent of Sap--Cohesion Supported by Tissue Pressure

Recent work contradicting both the assumptions of the CohesionTheory, and the tensions measured in the xylem sap by the pressure-chamber,is reviewed. Measurements with the xylem-pressure probe revealpressures in vessels around 0 bar absolute, and no detectablegradients of pressure with tree height. Under high water stress,pressures down to -6 bar were found, but then cavitations occurredvery readily. Also, measurements of the cavitation thresholdsof water show an average threshold of about -2 bar. The uncertainfoundations of the Cohesion Theory are recalled from the yearsbefore 1965. Soon after that date, Scholander's measurementswith the pressure chamber were agreed to have confirmed thetheory and the existence of high tensions in the xylem. Before1965, many experiments over many years pointed to the conclusionsnow rediscovered, viz., no high tensions, and no gradients oftension. A resolution of these paradoxes is offered in the formof a new theory. This proposes that the driving force and thetransmission of the force are the same as in the Cohesion Theory,but the operating pressure of the xylem is raised into a stablerange by compensating tissue pressures pressing upon the trachearyelements. The tissue pressure does not propel the transpirationstream, which is still driven by evaporation, but protects thestream from cavitation. Evidence is presented for the existenceof positive pressures in roots, wood, and leaves. It is shownthat the anatomy of roots, wood, and monocotyledon and cryptogamvascular bundles is organized so that pressure is confined bymechanical barriers, and exerted upon the tracheary elementsby the living cells of the phloem and the xylem parenchyma.The Compensating-Pressure Theory also explains, among otherthings, root pressure, the function of the endodermis, the structureof wood, the constant association of xylem and phloem, the absenceof gas spaces in vascular tissue, the absence of a gravitationalgradient in the xylem, bleeding from cut palm inflorescences,how insects are able to withdraw sap from the xylem, and thevariable that is measured by the pressure chamber. This instrumentmeasures the water potential, but this is the potential notof xylem in tension, but of the compensating pressure appliedto the xylem. The requirements of the Theory are explained,and a number of predictions are made which are open to experimentaltesting.Copyright 1995, 1999 Academic Press Ascent of sap, cavitation, cohesion theory, endodermis, pressure chamber, root pressure, stem pressure, tissue pressure, transpiration, water potential, wood anatomy, xylem pressure     

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.     

Mechanism of freeze-induced embolism in Fagus sylvatica L.

 The mechanism of freeze stress-induced embolism in Fagus sylvatica L. branches was analyzed under controlled conditions. Excised branches were exposed to successive freeze-thaw cycles in temperature controlled chambers. Thermocouples were placed on the bark to detect sap freezing exotherms. The degree of xylem embolism was estimated after each cycle by the loss of hydraulic conductivity. After one freeze-thaw cycle the degree of embolism was found to decrease with xylem specific hydraulic conductivity, small apical shoots being more susceptible to embolism. Exotherms revealed that distal shoots were freezing first and exuded sap as a result of water expansion. The lower water content in apical shoots upon freezing probably induced higher sap tensions which promoted air bubble expansion and vessel cavitation preferentially near the apices. When the decrease in water content was experimentally prevented, embolism developed to a lesser extent. The higher vulnerability of shoot apices may protect the rest of the branch from winter damage. Received: 29 May 1998 / Accepted: 15 August 1998     

Thermal dissipation probe measurements of sap flow in the xylem of trees documenting dynamic relations to variable transpiration given by instantaneous weather changes and the activities of a mistletoe xylem parasite

The thermal dissipation probe was described in the early 1930s for the demonstration of a volume and mass flow of sap in the conductive elements of the xylem in trees. It was subsequently developed further and is now widely used in physiological ecology including measurements in the field. Thermal dissipation demonstrates the occurrence of sap flow and allows determination of its velocity. Here we report simultaneous continuous measurements of sap flow using the thermal dissipation technique and of transpiration by infrared gas analysis for diurnal and annual cycles in a deciduous and an evergreen oak tree, Quercus robur L. and Quercus turneri Willd., respectively, in a deciduous and an evergreen conifer, Larix decidua Mill. and Pinus griffithii McClell., respectively, and the host/mistletoe consortium of the deciduous linden Tilia mandschurica Rupr. & Max. and the evergreen Viscum album L. We show (1) that in diurnal cycles sap flow closely follows dynamic changes of the rate of transpiration elicited by daily fluctuations of weather parameters (sunshine, cloudiness, air temperature and humidity), (2) that in annual cycles sap flow reflects autumnal yellowing and shedding of leaves of the deciduous trees. We report for the first time comparative measurements of sap flow towards mistletoe shoots and host branches in a parasite/host consortium. This demonstrates (3) that mistletoes maintain considerably larger sap flow rates in their xylem conduits than the adjacent host branches dragging the transpiration stream of their host towards their own shoots. We also show (4) that even after the deciduous host has shed its leaves and itself does not transpire any more the evergreen mistletoe towards its shoots can maintain the persistence of a continuous sap flow via the stem and branches of the host as long as frost does not prevent that. The work presented underlines the contention that transpiration is the driving force for sap flow with continuous files of water in the xylem. It shows for the first time that mistletoes direct the flow of water through host roots and stems towards its own shoots by not only performing stronger transpiration as it is known from the literature but also by maintaining larger sap flow rates in the xylem conduits of its stems.     

Excising stem samples underwater at native tension does not induce xylem cavitation

Xylem resistance to water stress‐induced cavitation is an important trait that is associated with drought tolerance of plants. The level of xylem cavitation experienced by a plant is often assessed as the percentage loss in conductivity (PLC) at different water potentials. Such measurements are constructed with samples that are excised underwater at native tensions. However, a recent study concluded that cutting conduits under significant tension induced cavitation, even when samples were held underwater during cutting. This resulted in artificially increased PLC because of what we have termed a ‘tension‐cutting artefact’. We tested the hypothesized tension‐cutting artefact on five species by measuring PLC at native tension compared with after xylem tensions had been relaxed. Our results did not support the tension‐cutting artefact hypothesis, as no differences were observed between native and relaxed samples in four of five species. In a fifth species (Laurus nobilis), differences between native and relaxed samples appear to be due to vessel refilling rather than a tension‐cutting effect. We avoided the tension‐cutting artefact by cutting samples to slightly longer than their measurement length and subsequent trimming of at least 0.5 cm of sample ends prior to measurement.     

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.     

Wood anatomy of some trees with diffuse- and ring-porous wood: Some functional and ecological interpretations

Abstract

The xylem conduit dimensions (i.e. their width and length) have been measured in 1-year-old internodes, nodes and node-to-petiole (N-P) junctions of three species with diffuse-porous wood, namely Ceratonia siliqua L., Laurus nobilis L. and Olea europaea L. as well as of three species with ring-porous wood, namely Quercus ilex L., Q. suber L. and Q. pubescens Willd‥ The xylem conduit diameter and length distributions have been related to the drought resistance strategies adopted by the six species. C. siliqua and Q. ilex (drought avoiding water spenders) showed the widest xylem conduits (each species within its characteristic pattern of wood anatomy). This is consistent with their high demand of efficient water transport to leaves. L. nobilis (drought avoiding water saver) showed relatively narrow xylem conduits, efficient enough, however, to assure water supply to leaves at the reduced transpiration rate exhibited by the species. O. europaea, Q. suber and Q. pubescens (drought tolerants) showed the narrowest xylem conduits but also the longest ones. The xylem system of C. siliqua and Q. ilex represented a good compromise between efficiency and safety of the water transport, the former as due to wide xylem conduits, the latter to the reduced xylem conduit length as well as to the strong «hydraulic constrictions» at their nodes and N-P junctions. The ecological interpretation of such hydraulic architecture is discussed.