Modeling xylem and phloem water flows in trees according to cohesion theory and Münch hypothesis

Water and solute flows in the coupled system of xylem and phloem were modeled together with predictions for xylem and whole stem diameter changes. With the model we could produce water circulation between xylem and phloem as presented by the Münch hypothesis. Viscosity was modeled as an explicit function of solute concentration and this was found to vary the resistance of the phloem sap flow by many orders of magnitude in the possible physiological range of sap concentrations. Also, the sensitivity of the predicted phloem translocation to changes in the boundary conditions and parameters such as sugar loading, transpiration, and hydraulic conductivity were studied. The system was found to be quite sensitive to the sugar-loading rate, as too high sugar concentration, (approximately 7 MPa) would cause phloem translocation to be irreversibly hindered and soon totally blocked due to accumulation of sugar at the top of the phloem and the consequent rise in the viscosity of the phloem sap. Too low sugar loading rate, on the other hand, would not induce a sufficient axial water pressure gradient. The model also revealed the existence of Münch “counter flow”, i.e., xylem water flow in the absence of transpiration resulting from water circulation between the xylem and phloem. Modeled diameter changes of the stem were found to be compatible with actual stem diameter measurements from earlier studies. The diurnal diameter variation of the whole stem was approximately 0.1 mm of which the xylem constituted approximately one-third.     

Sugar demand of ripening grape berries leads to recycling of surplus phloem water via the xylem

We tested the common assumption that fleshy fruits become dependent on phloem water supply because xylem inflow declines at the onset of ripening. Using two distinct grape genotypes exposed to drought stress, we found that a sink‐driven rise in phloem inflow at the beginning of ripening was sufficient to reverse drought‐induced berry shrinkage. Rewatering accelerated berry growth and sugar accumulation concurrently with leaf photosynthetic recovery. Interrupting phloem flow through the peduncle prevented the increase in berry growth after rewatering, but interrupting xylem flow did not. Nevertheless, xylem flow in ripening berries, but not berry size, remained responsive to root or shoot pressurization. A mass balance analysis on ripening berries sampled in the field suggested that phloem water inflow may exceed growth and transpiration water demands. Collecting apoplastic sap from ripening berries showed that osmotic pressure increased at distinct rates in berry vacuoles and apoplast. Our results indicate that the decrease in xylem inflow at the onset of ripening may be a consequence of the sink‐driven increase in phloem inflow. We propose a conceptual model in which surplus phloem water bypasses the fruit cells and partly evaporates from the berry surface and partly moves apoplastically to the xylem for outflow.     

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。     

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     

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     

A closed‐form solution for steady‐state coupled phloem/xylem flow using the Lambert‐W function

A closed‐form solution for steady‐state coupled phloem/xylem flow is presented. This incorporates the basic Münch flow model of phloem transport, the cohesion model of xylem flow, and local variation in the xylem water potential and lateral water flow along the transport pathway. Use of the Lambert‐W function allows this solution to be obtained under much more general and realistic conditions than has previously been possible. Variation in phloem resistance (i.e. viscosity) with solute concentration, and deviations from the Van't Hoff expression for osmotic potential are included. It is shown that the model predictions match those of the equilibrium solution of a numerical time‐dependent model based upon the same mechanistic assumptions. The effect of xylem flow upon phloem flow can readily be calculated, which has not been possible in any previous analytical model. It is also shown how this new analytical solution can handle multiple sources and sinks within a complex architecture, and can describe competition between sinks. The model provides new insights into Münch flow by explicitly including interactions with xylem flow and water potential in the closed‐form solution, and is expected to be useful as a component part of larger numerical models of entire plants.     

华北落叶松树体储水利用及其对土壤水分和潜在蒸散的响应: 基于模型模拟的分析

 树体储水在树木水分传输中具有重要的作用, 不仅为蒸腾提供水分来源, 还具有缓冲作用, 可防止木质部导管水势过低以至于水分传输的失败。树体储水动态及其利用的研究对于认识树木对水分胁迫的响应机制具有重要意义。该研究构建了包含树体储水释放-补充作用的树干水分传输模型, 可模拟计算林分小时尺度的冠层蒸腾、边材液流、树体储水与木质部导管水流交换过程, 并以六盘山北侧的华北落叶松(Larix principis-rupprechtii)人工林为例, 在林分水平分析树体储水利用及其 与土壤水分和潜在蒸散之间的关系。检验结果表明, 该模型能够精确地模拟出林分边材液流的日变化特征, 模拟与观测的小时液流速率决定系数R²为0.91 (n = 2 352)。模拟结果表明, 在典型晴朗天气下, 在日出时树体储水利用启动, 至9:00左右达到峰值(0.14 mm?h^–1), 午间降至0, 下午降为负值直至午夜, 即进入树体补水阶段; 树体储水日使用量(DJ_z)为0.04–0.58 mm?d^–1, 与日蒸腾量(DT_r)成正相关(R² = 0.91), 对蒸腾的贡献为25.6%。分析结果表明, 当潜在蒸散(ET_p)低于4.9 mm?d^–1时, ET_p是华北落叶松树体储水利用的主要驱动因子, DJ_z与ETp成正相关(R² = 0.68); 当ET_p高于4.9 mm?d^–1时, DJ_z随着ET_p的增加呈现降低趋势; DJ_z与土壤水势没有显著相关关系(p > 0.05), 但最大树体储水日使用量(DJ_zmax)与土壤水分含量成正相关(R² = 0.79), 说明土壤水分是树体储水利用的限制因子。     

Linking phloem function to structure: Analysis with a coupled xylem-phloem transport model

We carried out a theoretical analysis of phloem transport based on Münch hypothesis by developing a coupled xylem-phloem transport model. Results showed that the maximum sugar transport rate of the phloem was limited by solution viscosity and that transport requirements were strongly affected by prevailing xylem water potential. The minimum number of xylem and phloem conduits required to sustain transpiration and assimilation, respectively, were calculated. At its maximum sugar transport rate, the phloem functioned with a high turgor pressure difference between the sugar sources and sinks but the turgor pressure difference was reduced if additional parallel conduits were added or solute relays were introduced. Solute relays were shown to decrease the number of parallel sieve tubes needed for phloem transport, leading to a more uniform turgor pressure and allowing faster information transmission within the phloem. Because xylem water potential affected both xylem and phloem transport, the conductance of the two systems was found to be coupled such that large structural investments in the xylem reduced the need for investment in the phloem and vice versa.     

An analysis of the accumulation of water and dry matter in tomato fruit

Abstract Previously published data from tomato plants grown in nutrient solutions having one of three electrical conductivities (2, 12 and 17 mS cm^?1) were analysed. The rate of water import into the fruit, and the proportion of this conducted by the xylem stream were calculated from the daily rates of transpiration and the net accumulation of water and calcium. The rate of water import decreased as the conductivity of the nutrient solution rose, the maximum daily import rates in the third week after pollination being 3.2, 3.0 and 1.8 g fruit^?1 d^?1 for fruit grown at 2, 12 and 17 mS cm^?1, respectively. During fruit development, the proportion of water imported via the xylem fell from 8–15% to 1–2% at maturity. The principal source of water for tomato fruit growth was phloem sap. Based on the daily rates of net dry matter accumulation, respiration and phloem water import, the calculated dry matter concentration of the phloem sap declined from 7 to 3%, or from 12.5 to 7.8% during fruit development in low or high salinity, respectively. The similar dry matter accumulation of fruit grown at different salinities was due to changes in both volume and concentration of phloem sap. Potassium salts in tomato fruit were calculated lo have contributed –0.29, –0.48 and –0.58 MPa to total fruit osmotic potential in the 2, 12 and 17 mS cm^?1 treatments, respectively, which accounted for 38% or 49% of the measured total osmotic potential of the 2 mS cm^?1 or 17 mS cm^?1 treatments. The contribution of hexoses to total fruit osmotic potential in the young fruit was from about –0.1 to –0.2 MPa at all salinities. The osmotic potential of tomato fruit is regulated more by potassium salts than by hexoses.     

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.     

Time lags for xylem and stem diameter variations in a Scots pine tree

Diameter variations in the xylem and whole stem (i.e. over bark) stem of a Scots pine (Pinus sylvestris L.) tree were measured at four heights over a 23 d period at 5 min intervals. Cross‐correlation analysis was used to calculate time lags between the measurements. Xylem diameter measurements at the different heights had time lags varying from 10 to 50 min, measurements at the lower heights lagging behind the most. This result was in good agreement with the cohesion theory of transpiration. For the whole stem diameter measurements, the treetop lagged behind all other heights and the shortest lags were midway along the stem. Changes in whole stem diameter always lagged behind those of xylem stem diameter (30–110 min), and at all heights. The considerable differences in the behaviour of xylem and whole stem diameter support the Münch hypothesis of phloem flow. Time lags calculated separately for the shrinkage (morning) and swelling (afternoon) periods indicated shorter time lags during the swelling periods. The non‐destructive methods used show promise in the simultaneous study of flow dynamics of xylem and phloem in trees.     

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.

     

Assimilate transport in phloem sets conditions for leaf gas exchange

Carbon uptake and transpiration in plant leaves occurs through stomata that open and close. Stomatal action is usually considered a response to environmental driving factors. Here we show that leaf gas exchange is more strongly related to whole tree level transport of assimilates than previously thought, and that transport of assimilates is a restriction of stomatal opening comparable with hydraulic limitation. Assimilate transport in the phloem requires that osmotic pressure at phloem loading sites in leaves exceeds the drop in hydrostatic pressure that is due to transpiration. Assimilate transport thus competes with transpiration for water. Excess sugar loading, however, may block the assimilate transport because of viscosity build‐up in phloem sap. Therefore, for given conditions, there is a stomatal opening that maximizes phloem transport if we assume that sugar loading is proportional to photosynthetic rate. Here we show that such opening produces the observed behaviour of leaf gas exchange. Our approach connects stomatal regulation directly with sink activity, plant structure and soil water availability as they all influence assimilate transport. It produces similar behaviour as the optimal stomatal control approach, but does not require determination of marginal cost of water parameter.     

Analysis of Growth and Water Relations of Tomato Fruits in Relation to Air Vapor Pressure Deficit and Plant Fruit Load

The influence of air vapor pressure deficit (VPD) and plant fruit load on the expansion and water relations of young tomato fruits grown in a glasshouse were evaluated under summer Mediterranean conditions. The contributions of phloem, xylem and transpiration fluxes to the fruit volume increase were estimated at an hourly scale from the growth curves of intact, heat-girdled and detached fruits, measured using displacement transducers. High VPD conditions reduced the xylem influx and increased the fruit transpiration, but hardly affected the phloem influx. Net water accumulation and growth rate were reduced, and a xylem efflux even occurred during the warmest and driest hours of the day. Changes in xylem flux could be explained by variations in the gradient of water potential between stem and fruit, due to changes in stem water potential. Misting reduced air VPD and alleviated the reduction in fruit volume increase through an increase in xylem influx and a decrease in fruit transpiration. Under low fruit load, the competition for assimilates being likely reduced, the phloem flux to fruits increased, similarly to the xylem and transpiration fluxes, without any changes in the fruit water potential. However, different diurnal dynamics among treatments assume variable contributions of turgor and osmotic pressure in F3 and F6 fruits, and hypothetical short-term variations in the water potential gradient between stem and fruit, preventing xylem efflux in F3 fruits.     

Direct and indirect measurements of Phloem turgor pressure in white ash

Direct determinations and indirect calculations of phloem turgor pressure were compared in white ash (Fraxinus americana L.). Direct measurements of trunk phloem turgor were made using a modified Hammel-type phloem needle connected to a pressure transducer. Turgor at the site of the direct measurements was calculated from the osmotic potential of the phloem sap and from the water potential of the xylem. It was assumed that the water potentials of the phloem and xylem were close to equilibrium at any one trunk location, at least under certain conditions. The water potential of the xylem was determined from the osmotic potential of xylem sap and from the xylem tension of previously bagged leaves, measured with a pressure chamber. The xylem tension of bagged leaves on a branch adjacent to the site of the direct measurements was considered equivalent to the xylem tension of the trunk at that point. While both the direct and indirect measurements of phloem turgor showed clear diurnal changes, the directly measured pressures were consistently lower than the calculated values. It is not clear at present whether the discrepancy between the two values lies primarily in the calculated or in the measured pressures, and thus, the results from both methods as described here must be regarded as estimates of true phloem turgor.     

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.     

Effects of the hydraulic coupling between xylem and phloem on diurnal phloem diameter variation

Measurements of diurnal diameter variations of the xylem and phloem are a promising tool for studying plant hydraulics and xylem-phloem interactions in field conditions. However, both the theoretical framework and the experimental verification needed to interpret phloem diameter data are incomplete. In this study, we analytically evaluate the effects of changing the radial conductance between the xylem and the phloem on phloem diameter variations and test the theory using simple manipulation experiments. Our results show that phloem diameter variations are mainly caused by changes in the radial flow rate of water between the xylem and the phloem. Reducing the hydraulic conductance between these tissues decreases the amplitude of phloem diameter variation and increases the time lag between xylem and phloem diameter variation in a predictable manner. Variation in the amplitude and timing of diameter variations that cannot be explained by changes in the hydraulic conductance, could be related to changes in the osmotic concentration in the phloem.     

A non-invasive measurement of phloem and xylem water flow in castor bean seedlings by nuclear magnetic resonance microimaging

A flow-sensitive nuclear magnetic resonance (NMR) microimaging technique was applied to measure directly the in-vivo water flow in 6-d-old castor bean seedlings. The achieved in-plane resolution of the technique allowed discrimination between xylem and phloem water flow. Both the xylem- and the phloem-average flow velocities in the intact seedling could be quantified. Furthermore, the total conductive cross-sectional area of the xylem vessels and the phloem sieve elements could be determined using the non-invasive and non-destructive NMR microimaging technique. Hence, it was possible to calculate the in-vivo volume flow rates for both xylem and phloem water flow. Our non-destructive technique showed that previously used methods to measure phloem water flow affected the flow rate itself. In the intact seedlings we found values of 16.6 l·h^–1, two fold lower than those previously estimated from phloem exudation rates. Finally, our results demonstrate for the first time that water is internally circulated between phloem and xylem, and that water flow within the xylem is maintained by this internally circulated water, even in the absence of any significant transpiration or evaporation.Abbreviation NMR nuclear magnetic resonance     

How do salinity and water stress affect transport of water,assimilates and ions to tomato fruits?

The study was conducted in order to determine whether water stress affects the accumulation of dry matter in tomato fruits similarly to salinity, and whether the increase in fruit dry matter content is solely a result of the decrease in water content. Although the rate of water transport to tomato fruits decreased throughout the entire season in saline water irrigated plants, accumulation rates of dry matter increased significantly. Phloem water transport contributed 80–85% of the total water transport in the control and water-stressed plants, and over 90% under salinity. The concentration of organic compounds in the phloem sap was increased by 40% by salinity. The rate of ions transported via the xylem was also significantly increased by salinity, but their contribution to fruit osmotic adjustment was less. The rate of fruit transpiration was also markedly reduced by salinity. Water stress also decreased the rate of water transport to the tomato fruit and increased the rate of dry matter accumulation, but much less than salinity. The similar changes, 10–15%, indicate that the rise in dry matter accumulation was a result of the decrease in water transport. Other parameters such as fruit transpiration rates, phloem and xylem sap concentration, relative transport via phloem and xylem, solutes contributing to osmotic adjustment of fruits and leaves, were only slightly affected by water stress. The smaller response of these parameters to water stress as compared to salinity could not be attributed to milder stress intensity, as leaf water potential was found to be more negative. Measuring fruit growth of girdled trusses, in which phloem flow was inactive, and comparing it with ungirdled trusses validated the mechanistic model. The relative transport of girdled as compared to ungirdled fruits resembled the calculated values of xylem transport.     

Water transport properties of seven woody species from the semi-arid Mu Us Sandy Land,China

Maintenance of water transport is very important for plant growth and survival. We studied seven woody species that inhabit the semi-arid Mu Us Sandy Land, China, to understand their strategies for maintaining hydraulic function. We evaluated water transport properties, including cavitation resistance, hydraulic recovery, and water loss regulation by stomatal control, which are associated with xylem structural and leaf physiological traits. We also discussed the water-use characteristics of these species by comparing them with those of species in other regions. Species with tracheids had higher levels of xylem resistance to cavitation and a smaller midday transpiration rate than the other species studied. Although species with vessels were less resistant to cavitation, some recovered hydraulic conductivity within 12 h of rehydration. Species with xylem tracheids could maintain their hydraulic function through resistance to cavitation and by relaxing xylem tension. Although species with vessels had less resistant xylem, they could maintain hydraulic function through hydraulic recovery even when xylem dysfunction occurred. Additionally, the species studied here were less resistant to cavitation than species in semi-arid environments, and equally or less resistant than species in humid environments. Rather than allow hydraulic dysfunction due to drought-induced dehydration, they may develop water absorption systems to avoid or recover quickly from hydraulic dysfunction. Thus, not only stem cavitation resistance to drought but also stem–root coordination should be considered when selecting plants for the revegetation of arid regions.