Evidence for discontinuous water columns in the xylem conduit of tall birch trees

The continuity of the xylem water columns was studied on 17- to 23-m tall birch trees (trunk diameter about 23 cm; first branching above 10 m) all year round. Fifty-one trees were felled, and 5-cm thick slices or 2-m long boles were taken at regular, relatively short intervals over the entire height of the trees. The filling status of the vessels was determined by (i) xylem sap extraction from trunk and branch pieces (using the gas bubble-based jet-discharge method and centrifugation) and from trunk boles (using gravity discharge); (ii) ¹H nuclear magnetic resonance imaging of slice pieces; (iii) infusion experiments (dye, ⁸⁶Rb⁺, D₂O) on intact trees and cut branches; and (iv) xylem pressure measurements. This broad array of techniques disclosed no evidence for continuous water-filled columns, as postulated by the Cohesion–Tension theory, for root to apex directed mass transport. Except in early spring (during the xylem refilling phase) and after extremely heavy rainfall during the vegetation period, cohesive/mobile water was found predominantly at intermediate heights of the trunks but not at the base or towards the top of the tree. Similar results were obtained for branches. Furthermore, upper branches generally contained more cohesive/mobile water than lower branches. The results suggest that water lifting occurs by short-distance (capillary, osmotic and/or transpiration-bound) tension gradients as well as by mobilisation of water in the parenchymatic tissues and the heartwood, and by moisture uptake through lenticels.     

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.     

Water relations in silver birch during springtime: How is sap pressurised?

• Positive sap pressures are produced in the xylem of birch trees in boreal conditions during the time between the thawing of the soil and bud break. During this period, xylem embolisms accumulated during wintertime are refilled with water. The mechanism for xylem sap pressurization and its environmental drivers are not well known.
• We measured xylem sap flow, xylem sap pressure, xylem sap osmotic concentration, xylem and whole stem diameter changes, and stem and root non‐structural carbohydrate concentrations, along with meteorological conditions at two sites in Finland during and after the sap pressurisation period.
• The diurnal dynamics of xylem sap pressure and sap flow during the sap pressurisation period varied, but were more often opposite to the diurnal pattern after bud burst, i.e. sap pressure increased and sap flow rate mostly decreased when temperature increased. Net conversion of soluble sugars to starch in the stem and roots occurred during the sap pressurisation period. Xylem sap osmotic pressure was small in comparison to total sap pressure, and it did not follow changes in environmental conditions or tree water relations.
• Based on these findings, we suggest that xylem sap pressurisation and embolism refilling occur gradually over a few weeks through water transfer from parenchyma cells to xylem vessels during daytime, and then the parenchyma are refilled mostly during nighttime by water uptake from soil. Possible drivers for water transfer from parenchyma cells to vessels are discussed. Also the functioning of thermal dissipation probes in conditions of changing stem water content is discussed.

     

Gradients and dynamics of inner bark and needle osmotic potentials in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst)

Preconditions of phloem transport in conifers are relatively unknown. We studied the variation of needle and inner bark axial osmotic gradients and xylem water potential in Scots pine and Norway spruce by measuring needle and inner bark osmolality in saplings and mature trees over several periods within a growing season. The needle and inner bark osmolality was strongly related to xylem water potential in all studied trees. Sugar concentrations were measured in Scots pine, and they had similar dynamics to inner bark osmolality. The sucrose quantity remained fairly constant over time and position, whereas the other sugars exhibited a larger change with time and position. A small osmotic gradient existed from branch to stem base under pre‐dawn conditions, and the osmotic gradient between upper stem and stem base was close to zero. The turgor in branches was significantly driven by xylem water potential, and the turgor loss point in branches was relatively close to daily minimum needle water potentials typically reported for Scots pine. Our results imply that xylem water potential considerably impacts the turgor pressure gradient driving phloem transport and that gravitation has a relatively large role in phloem transport in the stems of mature Scots pine trees.     

Xylem Flow and its Driving Forces in a Tropical Liana: Concomitant Flow-Sensitive NMR Imaging and Pressure Probe Measurements

Abstract: Flow-sensitive NMR imaging and pressure probe techniques were used for measuring xylem water flow and its driving forces (i.e., xylem pressure as well as cell turgor and osmotic pressure gradients) in a tropical liana, Epipremnum aureum. Selection of tall specimens allowed continuous and simultaneous measurements of all parameters at various distances from the root under diurnally changing environmental conditions. Well hydrated plants exhibited exactly linearly correlated dynamic changes in xylem tension and flow velocity. Concomitant multiple-probe insertions along the plant shoot revealed xylem and turgor pressure gradients with changing magnitudes due to environmental changes and plant orientation (upright, apex-down, or horizontal). The data suggest that in upright and - to a lesser extent - in horizontal plants the transpirational water loss by the cells towards the apex during the day is not fully compensated by water uptake through the night. Thus, longitudinal cellular osmotic pressure gradients exist. Due to the tight hydraulic coupling of the xylem and the tissue cells these gradients represent (besides the transpiration-induced tension in the xylem) an additional tension component for anti-gravitational water movement from the roots through the vessels to the apex.     

Intact plant magnetic resonance imaging to study dynamics in long-distance sap flow and flow-conducting surface area

Due to the fragile pressure gradients present in the xylem and phloem, methods to study sap flow must be minimally invasive. Magnetic resonance imaging (MRI) meets this condition. A dedicated MRI method to study sap flow has been applied to quantify long-distance xylem flow and hydraulics in an intact cucumber (Cucumis sativus) plant. The accuracy of this MRI method to quantify sap flow and effective flow-conducting area is demonstrated by measuring the flow characteristics of the water in a virtual slice through the stem and comparing the results with water uptake data and microscopy. The in-plane image resolution of 120 x 120 microm was high enough to distinguish large individual xylem vessels. Cooling the roots of the plant severely inhibited water uptake by the roots and increased the hydraulic resistance of the plant stem. This increase is at least partially due to the formation of embolisms in the xylem vessels. Refilling the larger vessels seems to be a lengthy process. Refilling started in the night after root cooling and continued while neighboring vessels at a distance of not more than 0.4 mm transported an equal amount of water as before root cooling. Relative differences in volume flow in different vascular bundles suggest differences in xylem tension for different vascular bundles. The amount of data and detail that are presented for this single plant demonstrates new possibilities for using MRI in studying the dynamics of long-distance transport in plants.     

Diurnal changes in xylem pressure and mesophyll cell turgor pressure of the lianaTetrastigma voinierianum: The role of cell turgor in long-distance water transport

Summary Long-term xylem pressure measurements were performed on the lianaTetrastigma 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 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. Considering that the xylem pressure is determined by the turgor pressure (and vice versa), the xylem pressure of the liana could not drop to — in agreement with the experimental results — less than -0.4 MPa, because this pressure corresponds to zero turgor pressure.     

三角叶滨藜根吸水特点与其抗盐性的关系

采用压力室和冰点渗透压计测定了三角叶滨藜在不同浓度NaCl的根系环境溶液中根木质部的压力势和伤流液的渗透势,并利用原子吸收分光光度计测定了植株和伤流液以及环境溶液中Na 含量。结果表明:随着根环境溶液NaCl浓度的增加,三角叶滨藜植株和木质部伤流液中Na 含量虽呈上升趋势,但根系的过滤系数和体内Na 相对累积量逐渐降低,说明三角叶滨藜根细胞对盐分有很强的过滤作用;木质部伤流液的渗透势随着环境溶液渗透势的降低而降低,但根木质部溶液的水势则逐渐高出根外环境溶液的渗透势;表明三角叶滨藜能够利用较低的木质部负压来抵抗根外溶液的低渗透势而反渗透吸水,并利用根细胞对盐分的过滤作用来避免从环境摄取过量的盐分。     

A phloem-sap feeder mixes phloem and xylem sap to regulate osmotic potential

Phloem-sap feeders (Hemiptera) occasionally consume the dilute sap of xylem, a behaviour that has previously been associated with replenishing water balance following dehydration. However, a recent study reported that non-dehydrated aphids ingested xylem sap. Here, we tested the hypothesis that the consumption of xylem sap, which has a low osmolality, is a general response to osmotic stresses other than dehydration. Alate aphids were subjected to different treatments and subsequently transferred onto a plant, where electrical penetration graph (EPG) was used to estimate durations of passive phloem sap consumption and active sucking of xylem sap. The proportion of time aphids fed on xylem sap (i.e., time spent feeding on xylem sap/total time spent feeding on phloem plus xylem sap) was used as a proxy of the solute concentration of the uptake. The proportion of time alate aphids fed on xylem sap increased: (1) with the time spent imbibing an artificial diet containing a solution of sucrose, which is highly concentrated in phloem sap and is mainly responsible for the high osmotic potential of phloem sap; (2) with the osmotic potential of the artificial diet, when osmotic potential excess was not related to sucrose concentration; and (3) when aphids were deprived of primary symbionts, a condition previously shown to lead to a higher haemolymph osmotic potential. All our results converge to support the hypothesis that xylem sap consumption contributes to the regulation of the osmotic potential in phloem-sap feeders.     

What are the driving forces for water lifting in the xylem conduit?

After Renner had shown convincingly in 1925 that the transpirational water loss generates tensions larger than 0.1 MPa (i.e. negative pressures) in the xylem of cut leafy twigs the Cohesion Theory proposed by Böhm, Askenasy, Dixon and Joly at the end of the 19th century was immediately accepted by plant physiologists. Introduction of the pressure chamber technique by Scholander et al. in 1965 enforced the general belief that tension is the only driving force for water lifting although substantial criticism regarding the technique and/or the Cohesion Theory was published by several authors. As typical for scientific disciplines, the advent of minimal‐ and non‐invasive techniques in the last decade as well as the development of a new, reliable method for xylem sap sampling have challenged this view. Today, xylem pressure gradients, potentials, ion concentrations and volume flows as well as cell turgor pressure gradients can be monitored online in intact transpiring higher plants, and within a given physiological context by using the pressure probe technique and high‐resolution NMR imaging techniques, respectively. Application of the pressure probe technique to transpiring plants has shown that negative absolute pressures (down to ? 0.6 MPa) and pressure gradients can exist temporarily in the xylem conduit, but that the magnitude and (occasionally) direction of gradients contrasts frequently the belief that tension is the only driving force. This seems to be particularly the case for plants faced with problems of height, drought, freezing and salinity as well as with cavitation of the tensile water. Reviewing the current data base shows that other forces come into operation when exclusively tension fails to lift water against gravity due to environmental conditions. Possible candidates are longitudinal cellular and xylem osmotic pressure gradients, axial potential gradients in the vessels as well as gel‐ and gas bubble‐supported interfacial gradients. The multiforce theory overcomes the problem of the Cohesion Theory that life on earth depends on water being in a highly metastable state.     

树木树液上升机理研究进展

水分在植物体内的运输一直是很多植物生理生态学家所关注的一个重要问题。介绍了内聚力学说的基本假设和其存在争议,总结了近年来这一研究领域的几个热点问题,主要包括：（1）木质部栓塞及其恢复机理;（2）木质部压力探针和压力室法测定的木质部张力值不一致的现象及其可能原因;（3）补偿压学说;（4）不同界面层张力以及输水管道的毛细作用力、薄壁细胞膨压和木质部渗透压、逆向蒸腾等在树木汁液上升中的贡献;（5）最近发现的存在于木质部导管伴胞和韧皮部薄壁细胞等质膜中的水孔蛋白在植物水分运输中的调控作用等。这些方面在解释树木的树液上升中都起着重要的作用。     

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.     

盐胁迫对大豆根系木质部压力和Na+吸收的影响

取栽培大豆的水培幼苗为材料,用木质部压力探针和原子吸收分光光度计测定了盐胁迫条件下其根木质部压力和伤流液中Na~+含量的变化,以分析大豆抗盐吸水的机制.结果表明:在25～150 mmol/L NaCl的浓度范围内,随着盐胁迫强度的增加,大豆根木质部负压力的绝对值逐渐增大,但相对负压力和根的径向反射系数则逐渐减小;木质部伤流液中Na~+含量逐渐增加,但Na~+的相对含量则逐渐降低.同时,虽然根系吸水所需的木质部负压力(压力势)及根木质部伤流液的渗透势随着盐胁迫强度的增加都有所下降,但两者共同作用使木质部水势下降的幅度远远小于根外溶液水势(渗透势)下降的幅度,即随着根外溶液盐浓度的升高,根木质部溶液的总水势逐渐高出根外溶液的水势.上述结果说明,在盐胁迫下大豆可以利用相对小的木质部负压力逆水势梯度吸水,且通过避免对Na~+的过量吸收来适应盐胁迫环境.     

Xylem Sap from Actinidia chinensis: Apparent Differences in Sap Composition Arising from the Method of Collection

Large differences in composition were found between xylem sapcollected from Actinidia chinensis (Chinese gooseberry or kiwifruit) as bleeding sap and sap collected by vacuum extraction.A comparison of saps collected by the two methods has littlemeaning, however, unless the position on the plant from whichsap was collected and the prior treatment of the plant are specified.Furthermore the composition of bleeding sap changes rapidlywith time, probably because of marked gradients in concentrationof individual solutes in the xylem sap from the base to thetop of the plant. Contamination of vacuum-extracted sap by cellularcontents was shown to be insignificant. Sap collected as bleeding sap and by vacuum extraction are ofsomewhat different origins. It would be difficult to predictthe composition of bleeding sap simply from a knowledge of vacuum-extractedsap: it may be similarly unwise to predict the composition ofthe transpiration stream from that of vacuum extracted sap. Actinidia chinensis, kiwi fruit, Chinese gooseberry, xylem sap, bleeding sap, vacuum-extraction     

Thermodynamic analysis of the interaction of the xylem water and phloem sugar solution and its significance for the cohesion theory

The cohesion theory explains water transport in trees by the evaporation of water in the leaves (transpiration), which in turn generates the tension required for sap ascent, i.e. the flow of pure water from the soil through the root system and the non-living cells of the tree (xylem tracheids) up to the leaves. Only a small part of this water flow entering the leaves is used in photosynthesis to produce sugar solution, which is transported from the leaves through the living cells (phloem) to everywhere in the tree where it is needed and used. The phloem sieves are connected to the xylem tracheids by water transparent membranes, which means that the upflow of pure water and downflow of sugar solution interact with each other, causing the osmotic pressure in the sugar solution (Münch model). In this paper we analyse this interaction with a thermodynamic approach and we show that some open questions in the cohesion theory can then perhaps be better understood. For example, why under a quite high tension the water can flow in the xylem mostly without any notable cavitation, and how the suction force itself depends on the cavitation. Minimizing Gibbs energy of the system of xylem and phloem, we derive extended vapor pressure and osmotic pressure equations, which include gas bubbles in the xylem conduits as well as the cellulose-air-water interface term. With the aid of the vapor pressure equation derived here, we estimate the suction force that the cavitation controlled by the phloem sugar solution can generate at high moisture contents. We also estimate the suction force that the transpiration can generate by moisture gradient at low moisture contents. From the general osmotic pressure equation we derive an equation for calculating the degree of cavitation with different sugar solution concentrations and we show the conditions under which the cavitation in the xylem is totally avoided. Using recent field measurement results for a Scotch pine, the theory is demonstrated by showing its predictions for possible amounts of cavitation or embolism from morning hours to late afternoon.     

Measurement of turgor pressure and its gradient in the Phloem of oak

A direct method is described for measuring the pressure in secondary phloem sieve tubes of oak trees. One end of a 26-gauge stainless steel tube was shaped such that when it penetrated the outer bark and transected a few sieve elements, it was stopped by the xylem so that small openings in the end allowed phloem sap to enter the tube. The other end of the stainless tube (phloem needle) was joined to a long glass capillary sealed at its other end to form a manometer for measuring phloem sap pressure. A method for measuring the average osmotic and turgor pressures in cells of leaves is also described. Phloem turgor pressures varied greatly in a series of phloem punctures around the trunk at 1.5 and at 6.3 meters. The variation in turgor pressure was always greater than the variation in osmotic pressure. In a series of turgor pressures arranged in descending order, the values in a sequence for the upper level was usually a little (0-3 atm) larger than the values for the lower level. These results may suggest that translocation of assimilate is favored by a small turgor pressure gradient, but they do more to emphasize the complications in measuring gradients in an elastic low resistance distribution system composed of contiguous longitudinal conduits. The results also imply that the sieve tubes are inflated with assimilate fluid under high pressure which can readily move longitudinally and with less pressure drop than would be necessary if the sieve tubes were rigid.     

Applications of the compensating pressure theory of water transport

Some predictions of the recently proposed theory of long-distance water transport in plants (the Compensating Pressure Theory) have been verified experimentally in sunflower leaves. The xylem sap cavitates early in the day under quite small water stress, and the compensating pressure P (applied as the tissue pressure of turgid cells) pushes water into embolized vessels, refilling them during active transpiration. The water potential, as measured by the pressure chamber or psychrometer, is not a measure of the pressure in the xylem, but (as predicted by the theory) a measure of the compensating pressure P. As transpiration increases, P is increased to provide more rapid embolism repair. In many leaf petioles this increase in P is achieved by the hydrolysis of starch in the starch sheath to soluble sugars. At night P falls as starch is reformed. A hypothesis is proposed to explain these observations by pressure-driven reverse osmosis of water from the ground parenchyma of the petiole. Similar processes occur in roots and are manifested as root pressure. The theory requires a pump to transfer water from the soil into the root xylem. A mechanism is proposed by which this pump may function, in which the endodermis acts as a one-way valve and a pressure-confining barrier. Rays and xylem parenchyma of wood act like the xylem parenchyma of petioles and roots to repair embolisms in trees. The postulated root pump permits a re-appraisal of the work done by evaporation during transpiration, leading to the proposal that in tall trees there is no hydrostatic gradient to be overcome in lifting water. Some published observations are re-interpreted in terms of the theory: doubt is cast on the validity of measurements of hydraulic conductance of wood; vulnerability curves are found not to measure the cavitation threshold of water in the xylem, but the osmotic pressure of the xylem parenchyma; if measures of xylem pressure and of hydraulic conductance are both suspect, the accepted view of the hydraulic architecture of trees needs drastic revision; observations that xylem feeding insects feed faster as the water potential becomes more negative are in accord with the theory; tyloses, which have been shown to form in vessels especially vulnerable to cavitation, are seen as necessary for the maintenance of P, and to conserve the supplementary refilling water. Far from being a metastable system on the edge of disaster, the water transport system of the xylem is ultrastable: robust and self-sustaining in response to many kinds of stress.     

Phloem water relations and translocation

Satisfactory measurements of phloem water potential of trees can be obtained with the Richards and Ogata psychrometer and the vapor equilibration techniques, although corrections for loss of dry weight and for heating by respiration are required for the vapor equilibrium values. The psychrometer technique is the more satisfactory of the 2 because it requires less time for equilibration, less tissue, and less handling of tissue. Phloem water potential of a yellow-poplar tree followed a diurnal pattern quite similar to that of leaves, except that the values were higher (less negative) and changed less than in the leaves.

The psychrometer technique permits a different approach to the study of translocation in trees. Measurements of water potential of phloem discs followed by freezing of samples and determination of osmotic potential allows estimation of turgor pressure in various parts of trees as the difference between osmotic potential and total water potential. This technique was used in evaluating gradients in water potential, osmotic potential, and turgor pressure in red maple trees. The expected gradients in osmotic potential were observed in the phloem, osmotic potential of the cell sap increasing (sap becoming more dilute) down the trunk. However, values of water potential were such that a gradient in turgor pressure apparently did not exist at a time when rate of translocation was expected to be high. These results do not support the mass flow theory of translocation favored by many workers.

     

Levels of Sorbitol in Bleeding Sap and in Xylem Sap in Relation to Leaf Mass and Assimilate Demand in Apple Trees

Removal of fruits and treatment with SADH (succinic acid 2,2-dimethylhydrazide) were used to change the balance between sources and sinks for photosynthates in Malus domestica‘Golden Delicious’. Sorbitol and sugar content were measured in bleeding sap, in xylem sap prepared by suction, and in 80% methanol extracts of the roots. Concentration as well as total amount of sorbitol in bleeding sap sampled in July and September were lower in fruiting than in defruited trees, and so was the total amount of sorbitol in xylem sap from the trunk. SADH treatment tended to reduce the sorbitol content. Sorbitol in root extracts, expressed as percentage of methanol (80%) insoluble root dry matter, was highest in fruiting trees; but root dry matter was here less than half of that in defruited trees. Sorbitol content in xylem sap as well as sorbitol + sugar percentage of roots showed a distinct maximum in late winter followed by a heavy decrease during spring. It is suggested that sorbitol in xylem sap during the growth season represents a return transport from the roots, and that the level of sorbitol in this return transport reflects, to a certain degree, the ratio between leaf area and assimilate demand by the tree.     

Xylem Sap from Actinidia chinensis: Gradients in Sap Composition

In early spring, just prior to leaf break, there were substantialgradients in the concentration of many solutes in xylem sapcollected by vacuum extraction from shoots of different ages,from the branches, trunk and roots of mature Chinese gooseberryvines (Actinidia chinensis Planchon) For most nutrients therewas a progressive fall in concentration from youngest shootsdown to the roots and nitrate was the only nutrient to showa trend in the opposite direction These gradients in concentrationare probably sufficient to account for most of the changes observedin the composition of bleeding xylem sap during the first fewhours of bleeding after a branch is cut or wounded Actinidia chinensis, Chinese gooseberry, kiwifruit, xylem sap, nutrient gradients