No abstract

The resurrection plant Myrothamnus flabellifolia has the ability to recover from repeated prolonged and extreme desiccation cycles. During the dry state the inner walls of the xylem vessels seemed to be covered, at least partly, by a lipid film as shown by Sudan III and Nile Red staining. The lipid film apparently functioned as an ' internal cuticle ' which prevented the adjacent parenchyma ray cells from complete water loss. The hydrophobic nature of the inner xylem walls was supported by the finding that benzene ascended as rapidly as water in the xylem of dry Myrothamnus branches. On watering, numerous lipid bodies were found in the water-conducting vessels, presumably formed from the lipid film and\or from lipids excreted from the adjacent living cells into the vessels. The presence of lipid bodies within the vessels, as well as the hydrophobic properties of the inner xylem walls, could explain the finding that the xylem pressure of hydrated, well watered plants (measured both under laboratory and field conditions with the xylem pressure probe) never dropped below c. k0.3 MPa and that cavitation occurred frequently at low negative xylem pressure values (k0.05 to k0.15 MPa). The xylem pressure of M. flabellifolia responded rapidly and strongly to changes in relative humidity and temperature, but less obviously to changes in irradiance (which varied between 10 and c. 4000 µmol m −# s −" ). The morphological position of the stomata in the leaves could explain the extremely weak and slow response of the xylem pressure of this resurrection plant to illumination changes. Stomata were most abundant in the furrows, and were thus protected from direct sunlight. Simultaneous measurements of the cell turgor pressure in the leaf epidermal cells (made by using the cell turgor pressure probe) revealed that the xylem and the cell turgor pressure dropped in a ratio of 1 : 0.7 on changes in the environmental parameters, indicating a quite close hydraulic connection and, thus, water equilibrium between the xylem and cellular compartments. An increase in irradiance of c. 700 µmol m −# s −" resulted in a turgor pressure decrease from 0.63 to 0.48 MPa. Correspondingly, the cell osmotic pressure increased from 1.03 to 1.22 MPa. From these values and by assuming water equilibrium, the osmotic pressure of the xylem sap was estimated to be 0.25-0.4 MPa. This value seems to be fairly high but may, however, be explained by the reduction of the water volume within the vessels due to the floating lipid bodies.

Seasonal variations in osmolality and components of xylem sap in tall birch trees were determined using several techniques. Xylem sap was extracted from branch and trunk sections of 58 trees using the very rapid gas bubble-based jet-discharge method. The 5-cm long wood pieces were taken at short intervals over the entire tree height. The data show that large biphasic osmolality gradients temporarily exist within the conducting xylem conduits during leaf emergence (up to 272 mosmol x kg(-1) at the apex). These gradients (arising mainly from glucose and fructose) were clearly held within the xylem conduit as demonstrated by (1)H NMR imaging of intact twigs. Refilling experiments with benzene, sucrose infusion, electron and light microscopy, as well as (1)H NMR chemical shift microimaging provided evidence that the xylem of birch represents a compartment confined by solute-reflecting barriers (radial: lipid linings/lipid bodies; axial: presumably air-filled spaces). These features allow transformation of osmolality gradients into osmotic pressure gradients. Refilling of the xylem occurs by a dual mechanism: from the base (by root pressure) and from the top (by hydrostatic pressure generated by xylem-bound osmotic pressure). The generation of osmotic pressure gradients was accompanied by bleeding. Bleeding could be observed at a height of up to 21 m. Bleeding rates measured at a given height decreased exponentially with time. Evidence is presented that the driving force for bleeding is the weight of the static water columns above the bleeding point. The pressure exerted by the water columns and the bleeding volume depend on the water-filling status of (communicating) vessels.

The water supply to leaves of 25 to 60 m tall trees (including high-salinity-tolerant ones) was studied. The filling status of the xylem vessels was determined by xylem sap extraction (using jet-discharge, gravity-discharge, and centrifugation) and by (1)H nuclear magnetic resonance imaging of wood pieces. Simultaneously, pressure bomb experiments were performed along the entire trunk of the trees up to a height of 57 m. Clear-cut evidence was found that the balancing pressure (P(b)) values of leafy twigs were dictated by the ambient relative humidity rather than by height. Refilling of xylem vessels of apical leaves (branches) obviously mainly occurred via moisture uptake from the atmosphere. These findings could be traced back to the hydration and rehydration of mucilage layers on the leaf surfaces and/or of epistomatal mucilage plugs. Xylem vessels also contained mucilage. Mucilage formation was apparently enforced by water stress. The observed mucilage-based foliar water uptake and humidity dependency of the P(b) values are at variance with the cohesion-tension theory and with the hypothesis that P(b) measurements yield information about the relationships between xylem pressure gradients and height.

Summary. Long-term xylem pressure measurements were performed on the liana Tetrastigma voinierianum (grown in a tropical greenhouse) between heights of 1 m and 9.5 m during the summer and autumn seasons with the xylem pressure probe. Simultaneously, the light intensity, the temperature, and the relative humidity were recorded at the measuring points. Parallel to the xylem pressure measurements, the diurnal changes in the cell turgor and the osmotic pressure of leaf cells at heights of 1 m and 5 m (partly also at a height of 9.5 m) were recorded. The results showed that tensions (and height-varying tension gradients) developed during the day time in the vessels mainly due to an increase in the local light intensity (at a maximum 0.4 MPa). The decrease of the local xylem pressure from positive, subatmospheric or slightly above-atmospheric values (established during the night) to negative values after daybreak was associated with an almost ] : 1 decrease in the cell turgor pressure of the mesophyll cells (on average from about 0.4 to 0,5 MPa down to 0.08 MPa). Similarly, in the afternoon the increase of the xylem pressure towards more positive values correlated with an increase in the cell turgor pressure (ratio of about 1 : 1). The cell osmotic pressure remained nearly constant during the day and was about 0.75-0.85 MPa between 1 m and 9,5 m (within the limits of accuracy). These findings indicate that the turgor pressure primarily determines the corresponding pressure in the vessels (and vice versa) due to the tight hydraulic connection and thus due to the water equilibrium between both compartments. An increase in the transpiration rate (due to an increase in light intensity) results in very rapid establishment of a new equilibrium state by an equivalent decrease in the xylem and cell turgor pressure. From the xylem, cell turgor, and cell osmotic pressure data the osmotic pressure (or more accurately the water activity) of the xylem sap was calculated to be about 0.35-0.45 MPa; this value was apparently not subject to diurnal changes. Con-*Correspondence and reprints: Institut fiir Pflanzenphysiologie, Universit~t Salzburg, Hellbrunnerstral3e 34, A-5020 Salzburg, Austria. E-mail: friedrich.bentrup @ sbg.ac.at sidering that the xylem pressure is determined by the turgor pressure (and vice versa), the xylem pressure of the liana could not drop toin agreement with the experimental results -less than -0.4 MPa, because this pressure corresponds to zero turgot pressure.