Foliar water supply of tall trees: evidence for mucilage-facilitated moisture uptake from the atmosphere and the impact on pressure bomb measurements

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.     

Comparative measurements of the xylem pressure ofNicotiana plants by means of the pressure bomb and pressure probe

Determination of the pressure in the water-conducting vessels of intactNicotiana rustica L. plants showed that the pressure probe technique gave less-negative values than the Scholander-bomb method. Even though absolute values of the order of −0.1 MPa could be directly recorded in the xylem by means of the pressure probe, pressures between zero and atmospheric were also frequently found. The data obtained by the pressure probe for excised leaves showed that the Scholander bomb apparently did not read the actual tension in the xylem vessles ofNicotiana plants. The possibility that the pressure probe gave false readings was excluded by several experimental controls. In addition, cavitation and leaks either during the insertion of the microcapillary of the pressure probe, or else during the measurements were easily recognized when they occurred because of the sudden increase of the absolute xylem tension to that of water vapour or to atmospheric, respectively. Tension values of the same order could also be measured by means of the pressure probe in the xylem vessels of pieces of stem cut from leaves and roots under water and clamped at both ends. The magnitude of the absolute tension depended on the osmolarity of the bathing solution which was adjusted by addition of appropriate concentrations of polyethylene glycol. Partial and uniform pressurisation of plant tissues or organs, or of entire plants (by means of the Scholander bomb or of a hyperbaric chamber, respectively) and simultaneous recording of the xylem tension using the pressure probe showed that a 1∶1 response in xylem pressure only occurred under a few circumstances. A 1∶1 response required that the xylem vessels were in direct contact with an external water reservoir and/or that the tissue was (pre-)infiltrated with water. Corresponding pressure-probe measurements in isolated vascular bundles ofPlantago major L. orP. lanceolata L. plants attached to a Hepp-type osmometer indicated that the magnitude of the tension in the xylem vessels was determined by the external osmotic pressure of the reservoir. These and other experiments, as well as analysis of the data using classical thermodynamics, indicated that the turgor and the internal osmotic pressure of the accessory cells along the xylem vessels play an important role in the maintenance of a constant xylem tension. This conclusion is consistent with the cohesion theory. In agreement with the literature (P.E. Weatherley, 1976, Philos. Trans. R. Soc. London Ser. B23, 435–444; 1982, Encyclopedia of plant physiology, vol. 12B, 79-109), it was found that the tension in the xylem of intact plants under normal and elevated ambient pressure (as measured with the pressure probe) under quasi-stationary conditions was independent of the transpiration rate over a large range, indicating that the conductance of the flow path must be flow-dependent.     

Drying of mucilage causes water repellency in the rhizosphere of maize: measurements and modelling

Background and Aims

Although maize roots have been extensively studied, there is limited information on the effect of root exudates on the hydraulic properties of maize rhizosphere. Recent experiments suggested that the mucilaginous fraction of root exudates may cause water repellency of the rhizosphere. Our objectives were: 1) to investigate whether maize rhizosphere turns hydrophobic after drying and subsequent rewetting; 2) to test whether maize mucilage is hydrophobic; and 3) to find a quantitative relation between rhizosphere rewetting, particle size, soil matric potential and mucilage concentration.

Methods

Maize plants were grown in aluminum containers filled with a sandy soil. When the plants were 3-weeks-old, the soil was let dry and then it was irrigated. The soil water content during irrigation was imaged using neutron radiography. In a parallel experiment, ten maize plants were grown in sandy soil for 5 weeks. Mucilage was collected from young brace roots growing above the soil. Mucilage was placed on glass slides and let dry. The contact angle was measured with the sessile drop method for varying mucilage concentration. Additionally, capillary rise experiments were performed in soils of varying particle size mixed with maize mucilage. We then used a pore-network model in which mucilage was randomly distributed in a cubic lattice. The general idea was that rewetting of a pore is impeded when the concentration of mucilage on the pore surface (g cm^?2) is higher than a given threshold value. The threshold value depended on soil matric potential, pore radius and contract angle. Then, we randomly distributed mucilage in the pore network and we calculated the percolation of water across a cubic lattice for varying soil particle size, mucilage concentration and matric potential.

Results

Our results showed that: 1) the rhizosphere of maize stayed temporarily dry after irrigation; 2) mucilage became water repellent after drying. Mucilage contact angle increased with mucilage surface concentration (gram of dry mucilage per surface area); 3) Water could easily cross the rhizosphere when the mucilage concentration was below a given threshold. In contrast, above a critical mucilage concentration water could not flow through the rhizosphere. The critical mucilage concentration decreased with increasing particle size and decreasing matric potential.

Conclusions

These results show the importance of mucilage exudation for the water fluxes across the root-soil interface. Our percolation model predicts at what mucilage concentration the rhizosphere turns hydrophobic depending on soil texture and matric potential. Further studies are needed to extend these results to varying soil conditions and to upscale them to the entire root system.

     

Continuous measurements of water tensions in the xylem of trees based on the elastic properties of wood

According to the cohesion theory for the ascent of water in vascular plants, significant tensions should develop in the water columns of transpiring trees. These tensions cause small but detectable changes in the diameter of the xylem as a consequence of adhesive forces between water molecules and the inner xylem walls. The diurnal time course of tension in the water columns in the xylem of the trunk of mature Scots pine (Pinus sylvestris L.) was measured during the summer of 1995 by means of a displacement transducer mounted on a rigid steel frame. The apparent elastic modulus of Scots pine wood in the radial direction (E  ^′ _r ) was determined in the laboratory and then used to estimate tensions from the measured displacement. Laboratory measurements on logs indicated that only the sapwood contributed to dimensional changes of the xylem. Corrections for thermal expansion of the system were included. Water tensions fell by 0.19 MPa over the course of the day, when needle water potentials fell by 0.50 MPa. Such data are consistent with the cohesion theory, and with the view that the hydraulic resistances to flow in above- and below-ground plant parts are of similar magnitude. Received: 23 November 1996 / Accepted: 11 February 1997     

Vertical rooting patterns of mature Quercus trees growing on different soil types in northern Germany

At two different sites in northern Germany, the vertical distribution patterns of roots with diameters of up to 10 mm were investigated with the trench profile wall technique beneath mature trees of Quercus petraea (Matt.) Liebl. growing on clayey (cambi-stagnic gleysol) or sandy soil (dystric cambisol). The rooting patterns were related to soil bulk density, maximum plant-available soil water and soil chemistry, and were compared with the rooting pattern of Q. petraea growing on a silty soil on limestone (luvisol) and with that of Q. robur L. growing on clayey loam (stagnic cambisol). All three Q. petraea sites differed in their vertical rooting pattern, as was indicated by significant differences in the parameter which was calculated from the cumulative root fraction (Y) for the cumulative rooting depth (d; Y=1–^d). At the clayey site, a relatively large fraction of roots was found in deeper soil layers (highest value); whereas, at the silty site, the rooting pattern was most superficial (lowest value). No significant difference in the vertical rooting pattern was observed between Q. petraea and Q. robur growing on clayey soil. The soil-area related biomasses of living roots did not differ between the clayey and the sandy site of Q. petraea.Comparisons of the rooting patterns with soil water relations and soil chemistry lead to the hypothesis that under the climatic conditions of Central Europe, the vertical root distribution of Q. petraea is more influenced by the availability of nutrients, especially that of nitrogen, than by the amounts of plant-available soil water.     

Components of water potential estimated from xylem pressure measurements in five tree species

Summary Pressure volume curves were measured with a pressure bomb in leaves collected in the field from Ilex opaca, Acer rubrum, Liquidambar styraciflua, Liriodendron tulipifera and Cornus florida. Water potential components were calculated from the curves. The species differed in the relationships measured. In all species the trends from summer to fall were toward lower (more negative) osmotic potentials, lower matric potentials more rapid loss of turgor with increasing leaf water deficit, and the occurrence of incipient plasmolysis at lower values of leaf water deficit. Initial osmotic potentials ranged from-14.8 to-19.8 bars, similar to values reported in the literature for other mesophytic plants. These values, however, were much higher than those reported for halophytes and xerophytes. The fraction of leaf water which contributes to the osmotic potential ranged from 0.74 to 0.98 in this study. Values reported for other mesophytes and for halophytes and xerophytes all fall well within this range. Patterns of component water potentials are discussed in relation to potential growth rates and water flow in the total plant system.     

The expansion of maize root-cap mucilage during hydration. 3. Changes in water potential and water content

Root-cap mucilage from aerial nodal roots of maize has been found to have water potential values of −11 MPa or lower when air dried. The value approaches 0 MPa within 2 min of hydration in distilled water. In this time the expanding gel absorbs only about 0.3% of the water content of fully expanded mucilage. It is concluded that the root-cap mucilage per se has almost no capacity to retain water in the rhizosphere. Any function that it may play in the slowing of root desiccation would be indirect. For example, mucilage might decrease pore size between and within soil aggregates by pulling the particles together in a cycle of nocturnal efflux of water from the root surface, and diumal dyring during transpiration.     

The effect of pressure upon the solubility of oxygen in water: Implications of the deviation from the ideal gas law upon measurements of fluorescence quenching

The effect of pressure upon the solubility of oxygen in water compressed up to 500 atmospheres (atm) was directly measured. Pressurized aqueous solutions were allowed to equilibrate with a low pressure atmosphere (air, total pressure, 1 atm; oxygen partial pressure, 0.2 atm) across a gas-permeable, pressure-resistant Teflon membrane, and the equilibrium molar concentration of oxygen determined chemically. The solubility of oxygen in water decreased exponentially with pressure up to approximately 200 atm. At higher pressures, decreases in the molar volume of oxygen in solution resulted in a slight deviation from this initial trend. Implications of these results upon measurements of fluorescence quenching by oxygen under conditions of elevated pressure, a probe for studying the structure of macromolecules, are discussed.     

Fast spatiotemporal smoothing of calcium measurements in dendritic trees

We discuss methods for fast spatiotemporal smoothing of calcium signals in dendritic trees, given single-trial, spatially localized imaging data obtained via multi-photon microscopy. By analyzing the dynamics of calcium binding to probe molecules and the effects of the imaging procedure, we show that calcium concentration can be estimated up to an affine transformation, i.e., an additive and multiplicative constant. To obtain a full spatiotemporal estimate, we model calcium dynamics within the cell using a functional approach. The evolution of calcium concentration is represented through a smaller set of hidden variables that incorporate fast transients due to backpropagating action potentials (bAPs), or other forms of stimulation. Because of the resulting state space structure, inference can be done in linear time using forward-backward maximum-a-posteriori methods. Non-negativity constraints on the calcium concentration can also be incorporated using a log-barrier method that does not affect the computational scaling. Moreover, by exploiting the neuronal tree structure we show that the cost of the algorithm is also linear in the size of the dendritic tree, making the approach applicable to arbitrarily large trees. We apply this algorithm to data obtained from hippocampal CA1 pyramidal cells with experimentally evoked bAPs, some of which were paired with excitatory postsynaptic potentials (EPSPs). The algorithm recovers the timing of the bAPs and provides an estimate of the induced calcium transient throughout the tree. The proposed methods could be used to further understand the interplay between bAPs and EPSPs in synaptic strength modification. More generally, this approach allows us to infer the concentration on intracellular calcium across the dendritic tree from noisy observations at a discrete set of points in space.     

Systolic blood pressure and systolic hypertension in adolescence of atomic bomb survivors exposed in utero

Annual medical examinations were conducted during adolescence for the in utero clinical study sample subjects exposed prenatally to the atomic bombs in Hiroshima and Nagasaki. Systolic blood pressure and several anthropometric measurements were recorded during these examinations. For 1014 persons exposed in utero, two types of longitudinal analyses were performed, for a total of 7029 observations (6.93 observations per subject) of systolic blood pressure (continuous data) and systolic hypertension (binary data) for persons aged 9 to 19 years. Body mass index (BMI) and/or body weight were considered in the analyses as potential confounders. For the measurements of systolic blood pressure, the common dose effect was 2.09 mmHg per Gy and was significant (P = 0.017). The dose by trimester interaction was suggestive (P = 0.060). A significant radiation dose effect was found in the second trimester (P = 0.001), with an estimated 4.17 mmHg per Gy, but in the first and third trimesters, radiation dose effects were not significant (P > 0.50). For prevalence of systolic hypertension, the radiation dose effect was significant (P = 0.009); the odds ratio at 1 Gy was 2.23 [95% confidence interval (CI): 1.23, 4.04], and the dose by trimester interaction was not significant (P = 0.778). The dose response of systolic hypertension had no dose threshold, with a threshold point estimate of 0 Gy (95% CI: <0.0, 1.1 Gy). The dose response for systolic blood pressure was most pronounced in the second trimester, the most active organogenesis period for the organs relevant to blood pressure.     

Gas-exchange and sap flow measurements of Salix viminalis trees in short-rotation forest

Simultaneous field measurements of transpiration and sap flow were performed on short-rotation Salix viminalis trees ranging in diameter from 1.5 to 3.5 cm (2-year-old shoots on 8-year-old stumps). Transpiration was measured using an open-top ventilated chamber enclosing the whole foliage of a tree. Sap flow was measured using a tree-trunk heat balance (THB) technique with a constant temperature difference and variable heat input. Both the instantaneous and daily values of water flux measured by the two absolute techniques agreed well with a difference of up to about 5%. In July, the hourly transpiration reached a maximum of about 0.2 kg m^–2 (leaf area) or 0.45 kg tree^–1, whereas maximum daily integrals reached 4 kg tree^–1. The response of sap flow rate to abrupt flux change when inducing emboli by cutting-off the stem was very rapid: the registered signal dropped by 85% within 10 min for a specimen with a projected leaf area of 2 m². For S. viminalis trees, transpiration was linearly correlated with stem cross-sectional area and with leaf area.     

Differences in transpiration of Norway spruce drought stressed trees and trees well supplied with water

The paper focuses on the evaluation of transpiration as a physiological process, which is very sensitive to drought stress. Reactions of 25-year-old Norway spruce (Picea abies (L.) Karst.) trees to drought were examined during 2009 summer. Sap flow rate (SF), meteorological and soil characteristics were measured continually. Vapour pressure deficit of the air (VPD) and cumulative transpiration deficit (KTD) was calculated. During the second half of the vegetation period, the decrease in soil water content was observed and irrigation was applied to a group of spruce trees, while the second group was treated under natural soil drought. On the days, when the differences in transpiration between irrigated (IR) and non-irrigated (NIR) trees were significant (21 days), transpiration of NIR trees was only 23% of the transpiration of IR trees. We found significant differences in transpiration when the soil water content (SWC) of NIR variant at a depth of 5–15 cm ranged from 10.4 to 13.7%. Under both regimes of water availability, daily transpiration significantly responded to atmospheric conditions. However, the influence of all assessed meteorological parameters on SF of NIR trees was significantly lower than on IR tree. The dependency of transpiration on evaporative demands of atmosphere decreased with the decreasing soil moisture. Cumulative transpiration deficit of the stand during the entire evaluated period was 50.9 mm. The difference between the transpiration of the mean NIR tree and of the mean IR tree was 278.8 L over the assessed period of 47 days (5.9 L per day). The transpiration of NIR trees was 40.3% from the transpiration of IR trees during this period.     

Oxygen isotope ratios between soil water and stem water of trees in pot experiments

Since the oxygen isotopic ratio of water extracted from stems reflects that of water taken up by roots, the stem water isotope ratio can be used to analyze the source of water for plant growth. However, it is known that the fractionation of isotopes during evaporation from the surface soil increases the isotope ratio in soil water drastically. In this study, it was experimentally confirmed that the stem water of Elaeocarpus sylvestris vs. ellipticus Hara seedlings is not isotopically similar to the water source in the case where evaporation from the soil occurs actively. However, since water in these plant bodies was replaced in about 2 days in the pot experiments, the 2-day-averaged values of the soil water isotope ratio approached the stem water isotope ratio. Thus, time-course samplings of the soil and stems, and measurements of the replacement time of water in the plant body (water volume in plant/transpiration rate) are recommended for correct interpretation of the isotopic signature of soil water and stem water.     

Thermal properties of sapwood of fruit trees as affected by anatomy and water potential: errors in sap flux density measurements based on heat pulse methods

Key message

The accuracy of heat pulse methods is compromised both by changes in sapwood water content and by the methodology applied to determine its thermal properties.

Abstract

Sapwood thermal properties and water content (F _w) natural variations affect the accuracy of heat pulse sap flow methods, as they are typically set as constants or calculated during zero flow conditions. In a first experiment, a characterisation of both thermal properties and some of their determining anatomical and functional factors was conducted on several fruit tree species assessing the reliability of different methodologies. Besides that, a second experiment was carried out to evaluate the errors of heat pulse methods arising from ignoring F _w variations. To do so, desorption curves were constructed, allowing the substitution of F _w changes with those of water potential (Ψ). Results of the first experiment showed considerable differences between species in both the thermal, anatomical and functional properties of sapwood. Apart from that, discrepancies between the methods applied to determine thermal properties were also found and their implications for some heat pulse methods are discussed. The analysis conducted for the second experiment indicated that large errors in sap flux density (J) determinations might occur when daily and seasonal variations of Ψ (and hence of F _w) are disregarded. The extent of these errors was influenced by the species and heat pulse technique. Thus, the heat ratio and T _max were, respectively, the least and most vulnerable methods to errors in J determinations associated with changes in F _w.     

Stem water storage and diurnal patterns of water use in tropical forest canopy trees

Stem water storage capacity and diurnal patterns of water use were studied in five canopy trees of a seasonal tropical forest in Panama. Sap flow was measured simultaneously at the top and at the base of each tree using constant energy input thermal probes inserted in the sapwood. The daily stem storage capacity was calculated by comparing the diurnal patterns of basal and crown sap flow. The amount of water withdrawn from storage and subsequently replaced daily ranged from 4 kg d^–1 in a 0·20-m-diameter individual of Cecropia longipes to 54 kg d^–1 in a 1·02-m-diameter individual of Anacardium excelsum, representing 9–15% of the total daily water loss, respectively. Ficus insipida, Luehea seemannii and Spondias mombin had intermediate diurnal water storage capacities. Trees with greater storage capacity maintained maximum rates of transpiration for a substantially longer fraction of the day than trees with smaller water storage capacity. All five trees conformed to a common linear relationship between diurnal storage capacity and basal sapwood area, suggesting that this relationship was species-independent and size-specific for trees at the study site. According to this relationship there was an increment of 10 kg of diurnal water storage capacity for every 0·1 m² increase in basal sapwood area. The diurnal withdrawal of water from, and refill of, internal stores was a dynamic process, tightly coupled to fluctuations in environmental conditions. The variations in basal and crown sap flow were more synchronized after 1100 h when internal reserves were mostly depleted. Stem water storage may partially compensate for increases in axial hydraulic resistance with tree size and thus play an important role in regulating the water status of leaves exposed to the large diurnal variations in evaporative demand that occur in the upper canopy of seasonal lowland tropical forests.     

Predation pressure on the arboreal epiphytic herbivores of larch trees in southern England

Abstract. 1. Quantitative collections were made of arthropod predators from larch trees at Ranmore Common, Surrey, from August 1977 to September 1978.
2. Spiders were numerically the most important predators accounting for 64% of the total.
3. Antisera of the main larch dwelling, epiphyte herbivore orders, Collembola and Psocoptera, and the other common arboreal herbivore group at Ranmore, the Honioptera (principally Psyllidae) were prepared in rabbits.
4. Serological determinations of predator diets indicated that, with a few exceptions, the predators were taking approximately equal proportions of the three prey groups.
5. This apparent non-specificity of the predators is analysed in relation to both the numerical abundance and biomass of the three prey types during the study period.
6. These arboreal arthropod predators are selecting for large but rare prey items in preference to small and common ones.
7. These findings are discussed in relation to the population dynamics of arboreal epiphytic herbivores.