Hydraulic propagation of pressure along immature and mature xylem vessels of roots of Zea mays measured by pressure-probe techniques

Turgor pressure was measured in cortical cells and in xylem elements of excised roots and roots of intact plants of Zea mays L. by means of a cell pressure probe. Turgor of living and hence not fully differentiated late metaxylem (range 0.6–0.8 MPa) was consistently higher than turgor of cortical cells (range 0.4–0.6 MPa) at positions between 40 and 180 mm behind the root tip. Closer to the tip, no turgor difference between the cortex and the stele was measured. The turgor difference indicated that late-metaxylem elements may function as nutrient-storage compartments within the stele. Excised roots were attached to the root pressure probe to precisely manipulate the xylem water potential. Root excision did not affect turgor of cortical cells for at least 8 h. Using the cell pressure probe, the propagation of a hydrostatic pressure change effected by the root pressure probe was recorded in mature and immature xylem elements at various positions along the root. Within seconds, the pressure change propagated along both early and late metaxylems. The half-times of the kinetics, however, were about five times smaller for the early metaxylem, indicating they are likely the major pathway of longitudinal water flow. The hydraulic signal dissipated from the source of the pressure application (cut end of the root) to the tip of the root, presumably because of radial water movement along the root axis. The results demonstrate that the water status of the growth zone and other positions apical to 20 mm is mainly uncoupled from changes of the xylem water potential in the rest of the plant.Abbreviations and Symbols CPP cell pressure probe - EMX early metaxylem - LMX Late metaxylem - P_c cell turgor - P_r root pressure - RPP root pressure probe - t_1/2,c half-time of water exchange across a single cell - t_1/2 half-time of water exchange across multiple cells We thank Antony Matista for his expert assistance in the construction and modification of instruments. The work was supported by grant DCB8802033 from the National Science Foundation and grant 91-37100-6671 from USDA, and by the award of a Feodor Lynen-Fellowship from the Alexander von Humboldt-Foundation (Germany) to J.F.     

New evidence for large negative xylem pressures and their measurement by the pressure chamber method

Pressure probe measurements have been interpreted as showing that xylem pressures below c. –0.4 MPa do not exist and that pressure chamber measurements of lower negative pressures are invalid. We present new evidence supporting the pressure chamber technique and the existence of xylem pressures well below –0.4 MPa. We deduced xylem pressures in water-stressed stem xylem from the following experiment: (1) loss of hydraulic conductivity in hydrated stem xylem (xylem pressure = atmospheric pressure) was induced by forcing compressed air into intact xylem conduits; (2) loss of hydraulic conductivity from cavitation and embolism in dehydrating stems was measured, and (3) the xylem pressure in dehydrated stems was deduced as being equal and opposite to the air pressure causing the same loss of hydraulic conductivity in hydrated stems. Pressures determined in this way are only valid if cavitation was caused by air entering the xylem conduits (air-seeding). Deduced xylem pressure showed a one-to-one correspondence with pressure chamber measurements for 12 species (woody angiosperms and gymnosperms); data extended to c. –10 MPa. The same correspondence was obtained under field conditions in Betula occidentalis Hook., where pressure differences between air- and water-filled conduits were induced by a combination of in situ xylem water pressure and applied positive air pressure. It is difficult to explain these results if xylem pressures were above –0.4 MPa, if the pressure chamber was inaccurate, and if cavitation occurred by some mechanism other than air-seeding. A probable reason why the pressure probe does not register large negative pressures is that, just as cavitation within the probe limits its calibration to pressures above c. –0.5 MPa, cavitation limits its measurement range in situ.     

Water and solute transport along developing maize roots

Hydraulic and osmotic properties were measured along developing maize (Zea mays L.) roots at distances between 15 and 465 mm from the root tip to quantify the effects of changes in root structure on the radial and longitudinal movement of water and solutes (ions). Root development generated regions of different hydraulic and osmotic properties. Close to the root tip, passive solute permeability (root permeability coefficient, P_sr) was high and selectivity (root reflection coefficient, _sr) low, indicative of an imperfect semipermeable root structure. Within the apical 100–150 mm, P_sr decreased by an order of magnitude and _sr increased significantly. Root hydraulic conductivity (Lp_r) depended on the nature of the force (hydrostatic and osmotic). Osmotic Lp_r was smaller by an order of magnitude than hydrostatic Lp_r and decreased with increasing distance from the root tip. Throughout the root, responses in turgor of cortical cells and late metaxylem to step changes in xylem pressure applied to the base of excised roots were measured at high spatial resolution. The resulting profiles of radial and longitudinal propagation of pressure showed that the endodermis had become the major hydraulic barrier in older parts of the root, i.e. at distances from the apex ä 150 mm. Other than at the endodermis, no significant radial hydraulic resistance could be detected. The results permit a detailed analysis of the root's composite structure which is important for its function in collecting and translocating water and nutrients.Abbreviations and Symbols CPP cell pressure probe - IT root segments with intact tips; - Lp_r root hydraulic conductivity - Lp_rh hydrostatic hydraulic conductivity of root - Lp_ro osmotic hydraulic conductivity of root - P_app hydrostatic pressure applied to cut end of root - P_c cell turgor - P_{c, cor} turgor of cortical cell - P_c,xyl turgor of late metaxylem vessel - P_ro stationary root pressure - P_r0,seal stationary root pressure of sealed root segment - P_sr solute permeability coefficient of root - RPP root pressure probe - TR root segments with tip removed - _sr reflection coefficient of root Dedicated to Professor Andreas Sievers on the occasion of his retirement     

Water transport in barley roots

Radial transport of water in excised barley (Hordeum distichon, cv. Villa) roots was measured using a new method based on the pressure-probe technique. After attaching excised roots to the probe, root pressures of 0.9 to 2.9 bar were developed. They could be altered either by changing the root pressure artificially (with the aid of the probe) or by changing the osmotic pressure of the medium in order to induce water flows across the root. The hydraulic conductivity of the barley roots (per cm² of outer root surface) was obtained in different types of experiments (initial water flow, pressure relaxations, constant water flow) and was (0.3–4.3)·10^-7 cm s^-1 bar^-1. The hydraulic conductivity of the root was by an order of magnitude smaller than the hydraulic conductivity of the cell membranes of cortical and epidermal cells (0.8–2.2)·10^-6 cm s^-1 bar^-1. The half-times of water exchange of these cells was 1–21 s and two orders of magnitude smaller than that of entire excised roots (100–770 s). Their volumetric elastic modulus was 15–305 bar and increased with increasing turgor. Within the root cortex, turgor was independent of the position of the cell within a certain layer and turgor ranged between 3 and 5 bar. The large difference between the hydraulic conductivity of the root and that of the cell membranes indicates that there is substantial cell-to-cell (transcellular plus symplasmic) transport of water in the root. When it is assumed that 10–12 membrane layers (plasmalemma plus tonoplast) in the epidermis, cortex and endodermis form the hydraulic resistance to water flow, a value for the hydraulic conductivity of the root can be calculated which is similar to the measured value. This picture for water transport in the root contradicts current models which favour apoplasmic water transport in the cortex.     

NaCl Uptake in Roots of Zea mays Seedlings: Comparison of Root Pressure Probe and EDX Data

The extent by which salinity affects plant growth depends partlyon the ability of the plant to exclude NaCl. To study the uptakeof NaCl into excised roots of Zea mays L. cv. ‘Tanker’,two different techniques were applied. A root pressure probewas used to record steady state as well as transient valuesof root (xylem) pressure upon exposure of the root to mediacontaining NaCl and KCl as osmotic solutes. In treatments withNaCl, pressure/time responses of the root indicated a significantuptake of NaCl into the xylem. NaCl induced kinetics were completelyreversible when the NaCl solution was replaced by an isosmoticKCl solution. This indicated a passive movement of Na⁺-saltsacross the root cylinder. Root samples were taken at differenttimes of exposure to NaCl and prepared for X-ray microanalysis(EDX analysis). Radial profiles of ion concentrations (Na⁺,K⁺, Cl^–) were measured in cell vacuoles and xylem vesselsalong the root axis. Na⁺ appeared rapidly in mature xylem (earlymetaxylem) and living xylem (late metaxylem) before it was detectablein vacuoles of the root cortex. EDX results confirmed that thekinetics observed by the pressure probe technique correspondedmainly to an influx of Na⁺-salts into early metaxylem. In latemetaxylem, the uptake of Na⁺ was associated with a decline ofK⁺. The Na⁺/K⁺ exchange indicated a mechanism to reduce sodiumfrom the transpiration stream. Ion localization, ion transport, maize, root pressure, salinity, water relations, X-ray microanalysis, Zea mays     

Canny's compensating pressure theory fails a test

Canny's compensating pressure theory for water transport (American Journal of Botany 85: 897–909) has evolved from the premise that cavitation pressures are only a few tenths of a megapascal negative (approximately −0.3 MPa). In contradiction, “vulnerability curves” indicate that xylem pressures can drop below −3 MPa in some species without causing a loss of hydraulic conductivity. Canny claims these curves do not measure the limits to negative pressure by cavitation, but rather the limits to the compensating tissue pressure that otherwise quickly refills cavitated conduits. Compensating pressure is derived from the turgor pressure of the living cells in the tissue. To test this claim, we compared vulnerability curves of Betula nigra stems given three treatments: (1) living control, (2) killed in a microwave oven, and (3) perfused with a −1.5 MPa (10% w/w) mannitol solution. According to Canny's theory, the microwaved and mannitol curves should show cavitation and loss of conductance beginning at approximately −0.3 MPa because in both cases, the turgor pressure would be eliminated or substantially reduced compared to controls. We also tested the refilling capability of nonstressed stems where compensating pressure would be in full operation and compared this with dead stems with no compensating pressure. According to Canny's interpretation of vulnerability curves, the living stems should refill within 5 min. Results failed to support the compensating tissue theory because (a) all vulnerability curves were identical, reaching a −1.5 MPa threshold before substantial loss of conductance occurred, and (b) killed or living stems had equally slow refilling rates showing no significant increase in conductivity after 30 min. In consequence, the cohesion theory remains the most parsimonious explanation of xylem sap ascent in plants.     

Primary responses of root and leaf elongation to water deficits in the atmosphere and soil solution

Plants experience drought by a limitation of water supply andby enhanced transpiration. Both processes tend to decrease theplant's water potential, but affect growth responses in theroot and leaf differently. The evaluation of the underlyingmechanisms leads to a discussion of recent studies on biophysicalaspects of cell expansion at a cellular, tissue and organ level.Two processes enable roots to compensate rapidly effects ofwater deficits originating in the medium: (i) adjustment ofthe minimum pressure in cells required for expansion (yieldthreshold), and (ii) solute transport within the elongationzone. Limitations of root growth are discussed with respectto hydraulic, mechanical, and solute relations in the root elongationzone. It is argued that the variable nature of both the yieldthreshold and solute transport challenges the applicabilityof the Lockhart concept to determine growth-related parametersfrom steady conditions of turgor and growth. On a whole organlevel, the attenuation of xylem pressure along the root is importantfor the differential response of root and leaf growth. Experimentalevidence is presented for the hydraulic separation of the elongationzones, which is closely related to root development and functioning.The data obtained over the past few years have been used toextend mathematical models of growth and water transport inroots. Key words: Extension growth, hydraulic conductivity, root development (xylem, endodermis), transport (water and solute), turgor pressure, water stress, xylem pressure, Zea mays     

Cell water potential,osmotic potential,and turgor in the epidermis and mesophyll of transpiring leaves

Water potential, osmotic potential and turgor measurements obtained by using a cell pressure probe together with a nanoliter osmometer were compared with measurements obtained with an isopiestic psychrometer. Both types of measurements were conducted in the mature region of Tradescantia virginiana L. leaves under non-transpiring conditions in the dark, and gave similar values of all potentials. This finding indicates that the pressure probe and the osmometer provide accurate measurements of turgor, osmotic potentials and water potentials. Because the pressure probe does not require long equilibration times and can measure turgor of single cells in intact plants, the pressure probe together with the osmometer was used to determine in-situ cell water potentials, osmotic potentials and turgor of epidermal and mesophyll cells of transpiring leaves as functions of stomatal aperture and xylem water potential. When the xylem water potential was-0.1 MPa, the stomatal aperture was at its maximum, but turgor of both epidermal and mesophyll cells was relatively low. As the xylem water potential decreased, the stomatal aperture became gradually smaller, whereas turgor of both epidermal and mesophyll cells first increased and afterward decreased. Water potentials of the mesophyll cells were always lower than those of the epidermal cells. These findings indicate that evaporation of water is mainly occurring from mesophyll cells and that peristomatal transpiration could be less important than it has been proposed previously, although peristomatal transpiration may be directly related to regulation of turgor in the guard cells.     

An artificial osmotic cell: a model system for simulating osmotic processes and for studying phenomena of negative pressure in plants

Abstract An artificial osmotic cell has been constructed using reverse osmosis membranes. The cell consisted of a thin film of an osmotic solution (thickness: 100 to 200 μm) containing a non-permeating solute and was bounded between the membrane and the front plate of a pressure transducer which continuously recorded cell turgor. The membrane was supported by metal grids to withstand positive and negative pressures (P). At maximum, negative pressures of up to –0.7 MPa (absolute) could be created within the film on short-term and pressures of up to –0.3 MPa could be maintained without cavitation for several hours. As with living plant cells, the application of osmotic solutions of a non-permeating solute resulted in monophasic relaxations of turgor pressure from which the hydraulic conductivity of the membrane (Lp) and the elastic modulus of the cell (?) could be estimated. The application of solutions with permeating solutes resulted in biphasic pressure relaxation curves (as for living cells) from which the permeability (P_s) and reflection (σ_s) coefficients could be evaluated for the given membrane. Lp, P_s, and σ_s were independent of P and did not change upon transition from the positive to the negative range of pressure. It is concluded that the artificial cell could be used to simulate certain transport properties of living cells and to study phenomena of negative pressure as they occur in the xylem and, perhaps, also in living cells of higher plants.     

Xylem pressure response in maize roots subjected to osmotic stress: determination of radial reflection coefficients by use of the xylem pressure probe

The absolute pressure in conducting xylem vessels of roots of 2-week-old, slowly transpiring intact maize plants (bathed in nutrition medium) was determined to be +0·024 ± 0·044 MPa using the xylem pressure probe. When the roots were subjected to osmotic stress (NaCI, KCI or sucrose), the xylem pressure decreased immediately and became more negative. However, the response of xylem pressure to osmotic stress was considerably attenuated, indicating that the radial reflection coefficients, σ₁₃ of the maize root for these solutes were rather low (between 0·2 and 0·4 depending on the concentration of the osmoticum). The low values of a, may be caused (partly) by unstirred layer effects. In repeated osmoticum/nutrition regimes a complex pattern of changes in xylem pressure was observed which was apparently linked to the interplay between transpiration and (passive and/or active) solute loading of the xylem. These processes were not observed when the roots were subjected to osmotic stress after excision. In this case, a biphasic response was observed comparable to that found for excised roots using the root pressure probe.     

Diurnal variation in the radial reflection coefficient of intact maize roots determined with the xylem pressure probe

Xylem pressure and its relative response to the imposition ofan external osmotic stress (the so-called radial reflectioncoefficient) were recorded in roots of intact maize plants usingthe xylem pressure probe technique. Consecutive insertion oftwo probes into the same xylem vessel or into adjacent vesselsof intact roots of plants exposed to high light intensity andsalt stress under laboratory conditions showed that the xylemtension was not changed by vessel probing. It was also shownby using the double probe approach that the plants were capableof overcoming artificially induced leakages. This and otherevidence reported in the literature convincingly demonstratedthat the probe accurately reads xylem pressure and xylem pressureresponses to osmotic stress. Additional experiments were performedon plants grown in a greenhouse at a subtropical latitude. Underthese conditions the plants were exposed to strong diurnal fluctuationsin light intensity, relative humidity and temperature. The resultsshowed that the absolute xylem pressure in the roots of untreatedplants decreased with increasing transpiration rate from positivevalues in the early morning to negative values around noon (averagevalue –0.15 MPa; maximum negative value –0.57 MPa).As the day progressed and the transpiration rate decreased,xylem pressure increased again to positive values. Correspondingly,the radial reflection coefficient for NaCI increased from aboutzero in the early morning to about unity at noon when transpirationreached its highest value and decreased again to very low valuestowards the evening. The data raise questions concerning conclusionsabout the mechanism of water transport in intact roots drawnfrom the low radial reflection coefficients measured on excisedroots using the root pressure probe. Key words: Xylem pressure probe, osmotic stress, reflection coefficient, transpiration, diurnal changes     

Diurnal and seasonal variation in root xylem embolism in neotropical savanna woody species: impact on stomatal control of plant water status

Vulnerability to water-stress-induced embolism and variation in the degree of native embolism were measured in lateral roots of four co-occurring neotropical savanna tree species. Root embolism varied diurnally and seasonally. Late in the dry season, loss of root xylem conductivity reached 80% in the afternoon when root water potential (psi root) was about -2.6 MPa, and recovered to 25-40% loss of conductivity in the morning when psi root was about -1.0 MPa. Daily variation in psi root decreased, and root xylem vulnerability and capacitance increased with rooting depth. However, all species experienced seasonal minimum psi root close to complete hydraulic failure independent of their rooting depth or resistance to embolism. Predawn psi root was lower than psi soil when psi soil was relatively high (> -0.7 MPa) but became less negative than psi soil, later in the dry season, consistent with a transition from a disequilibrium between plant and soil psi induced by nocturnal transpiration to one induced by hydraulic redistribution of water from deeper soil layers. Shallow longitudinal root incisions external to the xylem prevented reversal of embolism overnight, suggesting that root mechanical integrity was necessary for recovery, consistent with the hypothesis that if embolism is a function of tension, refilling may be a function of internal pressure imbalances. All species shared a common relationship in which maximum daily stomatal conductance declined linearly with increasing afternoon loss of root conductivity over the course of the dry season. Daily embolism and refilling in roots is a common occurrence and thus may be an inherent component of a hydraulic signaling mechanism enabling stomata to maintain the integrity of the hydraulic pipeline in long-lived structures such as stems.     

Leaf illumination and root cooling inhibit bean leaf expansion by decreasing turgor pressure

Phaseolus vulgaris plants with expanding primary leaves weresubjected to dark-light or light-dark transition at a root temperatureof 25 °C, or to root cooling to 10 °C. Illuminationor darkening caused rapid changes in water flux through theplants and in epidermal turgor pressure when analysed by pressureprobe. However, these were not concurrent with variations inbulk leaf water potential and turgor pressure as determinedby the pressure chamber method. In addition, the turgor pressureof epidermis measured with the pressure probe was invariably0.05 to 0.15 MPa lower than that measured in bulk tissue withthe pressure chamber. Cooling roots to 10°C induced waterstress and wilting. Both techniques indicated a decrease ofturgor pressure, but a 20-30 min lag was observed with the pressurechamber. Due to stomatal closure and decreased transpiration,root-cooled plants regained cell turgor after 5-7 h of cooling,but bulk tissue and epidermal turgor (as well as leaf growthrate) remained significantly lower than control levels. Thesefindings indicate that changes in turgor pressure as the resultof hydraulic signalling are sufficient to explain the rapidchanges in growth rate following illumination or cooling reportedin earlier work (Sattin et al 1990). They also indicate thatdata obtained by use of the pressure chamber must be treatedwith caution. Key words: Phaseolus vulgaris, expansion growth, water relations, hydraulic signalling, pressure probe, pressure chamber     

胡杨木质部水分传导对盐胁迫的响应与适应北大核心CSCD

解析植物木质部导水率对逆境的响应和适应对促进植物抗逆性机理研究和受损植被恢复具有重要意义。该文以荒漠河岸林建群种胡杨(Populus euphratica)为研究对象,系统分析了胡杨幼株根、茎、叶水分传输通道对不同浓度盐胁迫的响应和适应。结果表明:(1)胡杨幼株根系对盐胁迫的敏感性高于茎和叶,盐胁迫下根系生长和根尖数显著受到抑制,根木质部易于发生栓塞,导水率明显降低。(2)胡杨幼株茎木质部导水率对盐胁迫的响应依盐浓度而定,轻度(0.05 mol·L–1 Na Cl)和中度(0.15 mol·L–1 Na Cl)盐胁迫下,胡杨可以通过协调导管输水的有效性和安全性来调节木质部的导水率,维持植物正常生长;重度(0.30 mol·L–1 Na Cl)盐胁迫下,胡杨茎木质部导管输水有效性和安全性均明显降低,木质部导水率显著下降,并伴随叶片气孔导度的显著降低,从而严重抑制了胡杨的光合和生长。     

Control of scion vigour by kiwifruit rootstocks is correlated with spring root pressure phenology

Root pressure was measured continuously over spring in eight clonal kiwifruit rootstocks selected from seven Actinidia species (A. chrysantha, A. deliciosa, A. eriantha, A. hemsleyana, A. kolomikta, A. macrosperma, A. polygama), using pressure transducers and miniature compression fittings. Rootstocks that promoted scion vigour developed root pressures up to 0.15 MPa before or during scion budburst, whereas those that reduced scion vigour developed root pressure up to 0.05 MPa only after scion shoot expansion. When several seasons were compared, the date of onset of root pressure and the magnitude of pressure achieved were consistent for each rootstock. Root pressure was first recorded between late July and early September in vigour-promoting rootstocks, while scion budburst and initial shoot growth were in late August and early September. Vigour-reducing rootstocks did not develop significant root pressure until October. The date of onset was similar for the grafted rootstock and ungrafted plant of the same clone, but was not clearly related to the timing of shoot growth by the ungrafted plant. In the grafted plants the leaf and xylem water potentials of the scion were more negative, midday turgor was 0.3-0.5 MPa lower, and wilting was sometimes observed in developing shoots growing on low-vigour rootstocks, indicating that water stress was contributing to reductions in growth. Leaf turgor was correlated with average root pressure but not pressure measured during the day, suggesting that root pressure was not supporting transpiration during peak flows and was, instead, indicative of higher root hydraulic conductance. The rapid temporal rise in root pressure observed each spring in the various rootstocks was not accompanied by changes in xylem sap solute potential, but when rootstock clones were compared those that developed higher root pressures had higher sap solute potentials. Xylem sap solute potential varied between rootstocks from -0.07 MPa to -0.15 MPa, while root pressures measured at the same time varied between 0.0 MPa and 0.09 MPa, suggesting that an osmotic mechanism could account for the observed root pressure. Differences in phenology between the rootstocks and scion appeared to account for the rootstock effects on shoot growth, and changes in root pressure provided a useful indication of seasonal changes in root hydraulic properties and solute transport behaviour.     

Xylem and cell turgor pressure probe measurements in intact roots of glycophytes: transpiration induces a change in the radial and cellular reflection coefficients

Xylem probe measurements in the roots of intact plants of wheat and barley revealed that the xylem pressure decreased rapidly when the roots were subjected to osmotic stress (NaCl or sucrose). The magnitude of the xylem pressure response and, in turn, that of the radial reflection coefficients (σ_r) depended on the transpiration rate. Under very low transpiration conditions (darkness and high relative humidity), σ_r assumed values of the order of about 0·2–0·4. The σ_r values of excised roots were also found to be rather low, in agreement with data obtained using the root pressure probe of Steudle. For transpiring plants (light intensities at least 10 μmol m^?2 s^?1; relative humidity 20–40%) the response was nearly 1:1, corresponding to radial reflection coefficients of σ_r= 1. Further increase of the light intensity to about 400 μmol m^?2 s^?1 resulted in a slight but significant decrease of the σ_r values to about 0·8. Similar measurements on maize roots confirmed our previous results (Zhu et al. 1995, Plant, Cell and Environment 18, 906–912) that, in intact transpiring plants at low light intensities of about 10 μmol m^?2 s^?1 and at relative humidities of 20–40% as well as in excised roots, the xylem pressure response was much less than expected from the external osmotic pressure (σ_r values 0·3–0·5). In contrast to wheat and barley, very high light intensities (about 700 μmol m^?2 s^?1) were needed to shift the radial reflection coefficients of maize roots to values of about 0·9. Osmotically induced xylem pressure changes were apparently linked to changes in turgor pressure in the root cortical parenchyma cells, as shown by simultaneous measurements of xylem and cell turgor pressure. In analogy to the σ_r values of the respective glycophytes, the σ_c values of the root cortical cells of wheat and barley were close to unity, whereas σ_c for maize was significantly smaller (about 0·7) under laboratory conditions. When the light intensity was increased up to about 700 μmol m^?2 s^?1 the cellular reflection coefficient of maize roots increased to about 0·95. In contrast to the σ_r values, the σ_c values of the three species investigated remained almost unchanged when the leaves were exposed to darkness and humidified air or when the roots were cut. The transpiration-dependent (species-specific) pattern of the cellular and radial reflection coefficients of the root compartment of the three glycophytes apparently resulted from (flow-dependent) concentration-polarization and sweep-away effects in the roots of intact plants. The data could be explained straightforwardly terms of theoretical considerations outlined previously by Dainty (1985, Acta Horticulturae 171, 21–31). The far-reaching consequences of this finding for root pressure probe measurements on excised roots, for the occurrence of pressure gradients under transpiring conditions, and for the non-linear flow-force relationships in roots found by other investigators are discussed.     

Hydraulic and osmotic properties of spruce roots

Hydraulic and osmotic properties of roots of 2-year-old Norwayspruce seedlings (Plcea abiea (L.) Karst) were investigatedusing different techniques (steady flow, pressure probe, andstop flow technique). Root pressures were measured using theroot pressure probe. Compared to roots of herbaceous plantsor deciduous trees, excised root systems of spruce did not developappreciable root pressure (-0.001 to 0.004 MPa or -10 to 40cm of water column). When hydrostatic pressure gradients wereused to drive water flows across the roots, hydraulic conductivities(Lp_r) were determined in two types of experiments: (i) rootpressure relaxations (using the root pressure probe) and (ii)steady flow experiments (pneumatic pressures applied to theroot system or xylem or partial vacuum applied to the xylem).Root Lp_r ranged between 0.2 and 810^–8m s^–1 MPa^–1(on average) depending on the conditions. In steady flow experiments,Lpr depended on the pressure applied (or on the flow acrossthe roots) and equalled (0.190.12) to (1.21.7)10^–8m s^–1 MPa^–1 at pressures between 0.2 and 0.4 MPaand (1.51.3)10^–8 m s^–1 MPa^–1 at appliedpressures between 0.8 and 1.0 MPa. When pressures or vacuumwere applied to the xylem, Lp_r values were similar. The hydraulicconductivity measured during pressure relaxations (transientwater flows) was similar to that obtained at high pressures(and water flows). Although there was a considerable scatterin the data, there was a tendency of the hydraulic conductivityof the roots to decrease with increasing size of the root system.When osmotic gradients were used to drive water flows, Lp_r valuesobtained with the root pressure probe were much smaller thanthose measured in the presence of hydrostatic gradients. Onaverage, a root Lp_r=0.01710^–8 was found for osmotic andLp_r=6.410^–8 m s^–1 MPa^–1 in correspondinghydrostatic experiments, i.e. the two values differed by a factorwhich was as large as 380. The same hydraulic conductivity asthat obtained in osmotic experiments using the pressure probewas obtained by the 'stop flow techniquel. In this technique,the suction created by an osmoticum applied to the root wasbalanced by a vacuum applied to the xylem. Lp_r values of rootsystems did not change significantly when measured for up to5 d. In osmotic experiments with different solutes (Na₂S0₄,K₂S0₄, Ca(NO₃)₂, mannitol), no passive uptake of solutes couldbe detected, i.e. the solute permeability was very low whichwas different from earlier findings on roots of herbs. Reflectioncoefficients of spruce roots (O were low for solutes for whichplant cell membranes exhibit values of virtually unity (     

Role of Pressure in Xylem Transport of Coconut and Other Palms

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

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.