High-pressure Flow Meter Research Articles

Use of a High-Pressure Flowmeter for Evaluating Hydraulic Characteristics of Plant Organs and Absorption Functions of Roots

The hydraulic characteristics of plant organs and the absorption functions of roots of corn and tomato plants were analyzed by using a specialized high-pressure flowmeter (HPFM), which was equipped with a root chamber as well as a coupling to connect to the plant organs. The HPFM can detect changes in the hydraulic properties of the respective parts of the plant body. The hydraulic resistances of the leaf and root decreased with an increase in the water flow rate; this dependence was remarkable at low water flow rates. On the other hand, the hydraulic resistance of the stem was only slightly affected by the water flow. However, a positive linear relationship between hydraulic conductance (reciprocal of resistance) and water flow rate was found for the shoot, stem, and root, respectively. Further, the rate of nutrient (NO3−) absorption by the root was evaluated from the water flow (root water absorption) and NO3− concentration in the xylem sap exuded from the root stump. The high-pressure flowmeter with the root chamber can be used for studying water and nutrient transport in plants and their responses to environmental elements.

Hydraulic efficiency and coordination with xylem resistance to cavitation, leaf function, and growth performance among eight unrelated Populus deltoides×Populus nigra hybrids

Tests were carried out to determine whether variations in the hydraulic architecture of eight Populus deltoides×Populus nigra genotypes could be related to variations in leaf function and growth performance. Measurements were performed in a coppice plantation on 1-year-old shoots under optimal irrigation. Hydraulic architecture was characterized through estimates of hydraulic efficiency (the ratio of conducting sapwood area to leaf area, A(X):A(L); leaf- and xylem-specific hydraulic conductance of defoliated shoots, k(SL) and k(SS), respectively; apparent whole-plant leaf-specific hydraulic conductance, k(plant)) and xylem safety (water potential inducing 50% loss in hydraulic conductance). The eight genotypes spanned a significant range of k(SL) from 2.63 kg s(-1) m(-2) MPa(-1) to 4.18 kg s(-1) m(-2) MPa(-1), variations being mostly driven by k(SS) rather than A(X):A(L). There was a strong trade-off between hydraulic efficiency and xylem safety. Values of k(SL) correlated positively with k(plant), indicating that high-pressure flowmeter (HPFM) measurements of stem hydraulic efficiency accurately reflected whole-plant water transport efficiency of field-grown plants at maximum transpiration rate. No clear relationship could be found between hydraulic efficiency and either net CO(2) assimilation rates, water-use efficiency estimates (intrinsic water-use efficiency and carbon isotope discrimination against (13)C), or stomatal characteristics (stomatal density and stomatal pore area index). Estimates of hydraulic efficiency were negatively associated with relative growth rate. This unusual pattern, combined with the trade-off observed between hydraulic efficiency and xylem safety, provides the rationale for the positive link already reported between relative growth rate and xylem safety among the same eight P. deltoides×P. nigra genotypes.

Comparison of Three Methods for Determination of Root Hydraulic Conductivity of Maize ( Zea mays L.) Root System

Three techniques of root pressure probe, pressure chamber and high pressure flow meter were used to measure the hydraulic conductivities ( Lp r ) of whole root systems of young maize ( Zea mays L.) seedlings grown hydroponically under either drought or normal water conditions. Compared to normal water conditions, drought stress simulated by polyethylene glycol 6 000 (osmotic potential = −0.2 MPa) reduced Lp r in the root system by over 50%. It indicated that water permeability in the roots decreased significantly when plants suffered from water shortages. Moreover, there was no significant difference ( P < 0.05) on the Lp r values in the root systems developed under a given water stress regime among the three techniques used. Therefore, all three methods are acceptable to study the hydraulic conductivity of maize seedling root systems. We have also highlighted some of the technical limitations of each method. It can be inferred that the root pressure probe is preferable for young maize seedlings because it is subtle and has the additional ability to determine solute transport properties, but the method is time consuming. Other advantages and disadvantages of each technique are discussed in order to acquaint researchers with basic information that could contribute to their choice of an appropriate technique for future studies.

Hydraulic connections of leaves and fruit to the parent plant in Capsicum frutescens (hot pepper) during fruit ripening.

The hydraulic architecture and water relations of fruits and leaves of Capsicum frutescens were measured before and during the fruiting phase in order to estimate the eventual impact of xylem cavitation and embolism on the hydraulic isolation of fruits and leaves before maturation/abscission. Measurements were performed at three different growth stages: (1) actively growing plants with some flowers before anthesis (GS1), (2) plants with about 50 % fully expanded leaves and immature fruits (GS2) and (3) plants with mature fruits and senescing basal leaves (GS3). Leaf conductance to water vapour as well as leaf and fruit water potential were measured. Hydraulic measurements were made using both the high-pressure flow meter (HPFM) and the vacuum chamber (VC) technique. The hydraulic architecture of hot pepper plants during the fruiting phase was clearly addressed to favour water supply to growing fruits. Hydraulic measurements revealed that leaves of GS1 plants as well as leaves and fruit peduncles of GS2 plants were free from significant xylem embolism. Substantial increases in leaf petiole and fruit peduncle resistivity were recorded in GS3 plants irrespective of the hydraulic technique used. The higher fraction of resistivity measured using the VC technique compared with the HPFM technique was apparently due to conduit embolism. The present study is the first to look at the hydraulics of leaves and fruits during growth and maturation through direct, simultaneous measurements of water status and xylem efficiency of both plant regions at different hours of the day.

Within plant resistance to water flow in tomato and sweet melons

Efficient water resource management in relation to water use and crop yields is premised on the knowledge of plant resistance to water flow. However, such studies are limited and for most crops, the within plant resistance to water flow remains largely unknown. In this study, within plant resistance to water transport (hydraulic conductance) was monitored in tomato (Lycopersicum esculuntum) and sweet melon (Citrullus lanatus) using the high pressure flow meter (HPFM) and evaporative flux (EF) methods. In the evaporative flux method, measurements of transpiration flux and leaf water potential were used to calculate the total resistance to water flow using Ohm’s law analogy. Measurements of tranpiration flux (Q) relationship, plant resistance calculated from the slope of their relationship, ranged from 6.57x10-01 to 2.27x10-03 Mpa m-2s-1 for tomato and sweet melon, respectively.The magnitude of whole plant hydraulic conductance calculated by the evaporative flux method and measured on the HPFM were not significantly different. This is probably due to that fact that Kp includes the hydraulic conductance of the root system, which offers the highest resistance to water flow in a plant, and the frictional resistance of the proximal part of the crown. Day time course of water relation parameters were monitored in melon and tomato (predawn, 1100 to 1400 h). The effect of time of day was pronounced on the dynamics of water relations, particularly around mid-day when sharp differences in the magnitudes of the measured water relation parameters were obtained. Minimum (predawn) leaf water potential (øl) ranged from 0.135 to 0.207 MPa for both crops. The greatest conductance was found in the leaf and the lowest in the root in both crops. While the highest within plant resistance to the flow is contained in the root system, the stemcomponent constitutes the least resistance (greatest conductance) to within plant flow of water in tomato. Repeated measurement analysis showed the existence of significant species effects on plant water relations (leaf water potential, evaporative/transpiration flux, xylem hydraulic conductance). Superiority within plant xylem transport and evaporative losses were obtained in tomato, from the relations of EF and Kh; this crop showed a higher hydraulic sufficiency than melon. The expression of hydraulic conductance of the root and shoot systemrelative to plant attributes did not eliminate differences in the magnitudes of conductance elements in tomato and melon. Differences obtained between melon and tomato in whole plant leaf and stem area specific hydraulic conductance (KI) indicate the carbon efficiency and, hence, the cost of resource allocation to areas of root surfaceand leaves for enhanced scavenging for water and mineral nutrients. Thus, application of mineral nutrients to enhance crop productivity should take into consideration the hydraulic conductance for specific plants.

Effects of water storage in the stele on measurements of the hydraulics of young roots of corn and barley

In standard techniques (root pressure probe or high-pressure flowmeter), the hydraulic conductivity of roots is calculated from transients of root pressure using responses following step changes in volume or pressure, which may be affected by a storage of water in the stele. Storage effects were examined using both experimental data of root pressure relaxations and clamps and a physical capacity model. Young roots of corn and barley were treated as a three-compartment system, comprising a serial arrangement of xylem/probe, stele and outside medium/cortex. The hydraulic conductivities of the endodermis and of xylem vessels were derived from experimental data. The lower limit of the storage capacity of stelar tissue was caused by the compressibility of water. This was subsequently increased to account for realistic storage capacities of the stele. When root water storage was varied over up to five orders of magnitude, the results of simulations showed that storage effects could not explain the experimental data, suggesting a major contribution of effects other than water storage. It is concluded that initial water flows may be used to measure root hydraulic conductivity provided that the volumes of water used are much larger than the volumes stored.

Aquaporin gene expression and apoplastic water flow in bur oak (Quercus macrocarpa) leaves in relation to the light response of leaf hydraulic conductance.

It has previously been shown that hydraulic conductance in bur oak leaves (Quercus macrocarpa Michx.), measured with the high pressure flow meter technique (HPFM), can significantly increase within 30 min following exposure to high irradiance. The present study investigated whether this increase could be explained by an increase in the cell-to-cell pathway and whether the response is linked to changes in the transcript level corresponding to aquaporin genes. Four cDNA sequences showing high similarity to members of the aquaporin gene family from other plant species were characterized from bur oak leaves and the expression levels of these cDNA sequences were examined in leaves by quantitative real-time PCR (QRT-PCR). No change was found in the relative transcript abundance corresponding to these four putative aquaporin genes in leaves with light-induced high hydraulic conductance (exposed to high irradiance) compared to leaves with low hydraulic conductance (exposed to low irradiance). However, in sun leaves that were exposed to different light levels prior to leaf collection (full sunlight, shade, and covered with aluminium foil for 16 h), the relative transcript levels of two of the putative aquaporin genes increased several-fold in shaded leaves compared to the sun-exposed or covered leaves. When the leaves were pressure-infiltrated with the apoplastic tracer dye trisodium 3-hydroxy-5,8,10-pyrenetrisulphonate (PTS3, 0.02%), there was no change in the PTS3 concentration of leaf exudates collected in ambient light or in high irradiance, but there was a small apoplastic acidification. There was also no change in PTS3 concentration between the leaves infiltrated under high irradiance with 0.02% PTS3 or with 0.1 mM HgCl2 in 0.02% PTS3. The results suggest that the putative aquaporin genes that were identified in the present study probably do not play a role in the light responses of hydraulic conductance at the transcript level, but they may function in regulating water homeostasis in leaves adapted to different light conditions. In addition, it is shown that high irradiance induced changes in the pH of the apoplast and that there does not appear to be a significant shift to the cell-to-cell mediated water transport in bur oak leaves exposed to high irradiance as measured by the apoplastic tracer dye.

The rapid light response of leaf hydraulic conductance: new evidence from two experimental methods

Previous studies have shown a rapid enhancement in leaf hydraulic conductance (K(leaf)) from low to high irradiance (from <10 to >1000 micromol photons m(-2) s(-1)), using the high-pressure flow meter (HPFM), for 7 of 14 tested woody species. However, theoretical suggestions have been made that this response might arise as an artifact of the HPFM. We tested the K(leaf) light response for six evergreen species using refined versions of the rehydration kinetics method (RKM) and the evaporative flux method (EFM). We found new evidence for a rapid, 60% to 100% increase in K(leaf) from low to high irradiance for three species. In the RKM, the leaf rehydration time constant declined by up to 70% under high irradiance relative to darkness. In the EFM, under higher irradiance, the flow rate increased disproportionately to the water potential gradient. Combining our data with those of previous studies, we found that heterobaric species, i.e. those with bundle sheath extensions (BSEs) showed a twofold greater K(leaf) light response on average than homobaric species, i.e. those without BSEs. We suggest further research to characterize this substantial dynamic at the nexus of plant light- and water-relations.

Root hydraulic conductivity measured by pressure clamp is substantially affected by internal unstirred layers

Using the root pressure probe in the pressure clamping (PC) mode, the impact of internal unstirred layers (USLs) was quantified for young corn roots, both in experiments and in computer simulations applying the convection/diffusion model of Knipfer et al. In the experiments, water flows (JVrs) during PC were analysed in great detail, showing that JVrs (and the apparent root hydraulic conductivity) were high during early stages of PC and declined rapidly during the first 80 s of clamping to a steady-state value of 40–30% of the original. The comparison of experimental results with simulations showed that, during PC, internal USLs at the inner surface of the endodermis substantially modify the overall force driving the water. As a consequence, JVr and Lpr were inhibited. Effects of internal USLs were minimized when using the pressure relaxation mode, when internal USLs had not yet developed. Additional stop-clamp experiments and experiments where the endodermis was punctured to reduce the effect of internal USLs verified the existence of internal USLs during PC. Data indicated that the role of pressure propagation along the root xylem for both PC and pressure relaxation modes should be small, as should the effects of filling of the capacities during root pressure probe experiments, which are discussed as an alternative model. The results supported the idea that concentration polarization effects at the endodermis (internal USLs) cause a serious problem whenever relatively large amounts of water (xylem sap) are radially moved across the root, such as during PC or when using the high-pressure flow meter technique.

Direct measurement of hydraulic properties in developing berries of Vitis vinifera L. cv Shiraz and Chardonnay

Berries of Vitis vinifera L. cv Shiraz can undergo weight loss during later stages of ripening. Existing published views on how weight loss occurs are based on changes in capacity of the vascular system to import water during development (McCarthy and Coombe, Australian Journal of Grape and Wine Research, 5, 17–21, 1999). One important element of these views is the proposed cessation of water flow through the xylem after veraison. We have now measured the water flow into berries of Shiraz and Chardonnay as they develop using the pressure probe and the high pressure flow meter (HPFM). The pressure probe connected to the pedicel of individual berries provided measurements of single berry hydraulic conductance. By systematic excision of tissue segments of the berry and pedicel we determined where in the pathway hydraulic conductance changed during development. The HPFM was used on whole bunches showing that berries (including pedicels) represent parallel high hydraulic resistances and that the hydraulic resistance of the bunch axis was rather small. The hydraulic conductance per berry could be determined from excision experiments. There was close agreement between the pressure probe and HPFM measurements. Both showed a ten-fold reduction in hydraulic conductance of whole berries from veraison to full ripeness. Shiraz had hydraulic conductances that were 2- to 5-fold higher than those for Chardonnay. Shiraz maintained a higher hydraulic conductance past 90 days after flowering than Chardonnay. The decrease in hydraulic conductance occurred in both the distal and proximal parts of the berry for both varieties. The pressure probe also provided measurements of the xylem pressure that non-transpiring berries could develop. These pressures were –0.2 to –0.1 MPa until veraison and increased to zero when the juice osmotic potential reached about –3 MPa in Chardonnay and –4 MPa in Shiraz. The results suggest values of the reflection coefficient of the osmotic barrier around the xylem vessels of about 0.1–0.2 at veraison decreasing to 0 at harvest. It is suggested that in addition to changes in xylem anatomy, aquaporins in berry membranes may play a role in regulating hydraulic conductance. Water movement from the berry back to the parent vine via the xylem (backflow) may be an important component of berry weight loss in Shiraz, particularly if the phloem ceases functioning at high osmotic potentials near maximum weight. Backflow could account for a weight loss of 43 mg per day in Shiraz berries for a relatively small gradient of 0.1 MPa.

Hydraulic resistance components of mature apple trees on rootstocks of different vigours

Dwarfing of fruit trees is often achieved through the use of dwarfing rootstocks. Dwarf trees are characterized by sustained reductions in vegetative growth during the lifetime of the tree. The dwarfing mechanism is not well understood, but it has been hypothesized that hydraulic properties of the rootstock and the graft union are involved. It is hypothesized here that leaf- or stem-specific resistance of at least one hydraulic component of the water transport system would be negatively correlated with rootstock 'vigour', and this could be useful for selection of rootstocks. Hydraulic resistance (R) of fully grown apple trees on a variety of rootstocks of different 'vigours' was measured. Most measurements were with the evaporative flux (EF) method, where water uptake measured with sap flow sensors was related to the pressure gradient from soil (taken as pre-dawn leaf) and midday root (taken as covered root-sucker), stem (from covered leaf), and exposed and shaded leaf water potentials (Psi(l)). R of trees on dwarfing M9 rootstock was compared with that of more vigorous MM106 and MM111 rootstocks in Israel and Vermont, USA. In Israel, M9 consistently had higher leaf-specific hydraulic resistance (R(l)) in the soil to scion stem pathway, but this difference was only significant for one summer. R was larger in M9 between the root and stem, implicating the graft union as the site of increased resistance. In Vermont, R(l) of 9- and 10-year-old trees on six rootstocks of various vigours was not consistently related to vigour, and stem-specific resistance (R(s)) increased with increasing vigour. High pressure flow meter (HPFM) measurements gave a lower R than the EF method in all but one case, perhaps indicating a significant amount of xylem dysfunction in these trees, and demonstrated the increased resistivity of stem sections that included dwarf graft unions as compared with non-graft stem sections. It is concluded that stem- and leaf-specific R are not consistently positively correlated with dwarfing, although the increased resistivity of the graft union in dwarfing rootstocks may influence the transport of water and other elements across the graft union, and therefore be involved in the dwarfing mechanism.

During measurements of root hydraulics with pressure probes, the contribution of unstirred layers is minimized in the pressure relaxation mode: comparison with pressure clamp and high-pressure flowmeter

The effects of unstirred layers (USLs) at the endodermis of roots of young maize plants (Zea mays L.) were quantified, when measuring the water permeability of roots using a root pressure probe (RPP) in the pressure relaxation (PR) and pressure clamp (PC) modes. Different from PRs, PCs were performed by applying a constant pressure for certain periods of time. Experimental data were compared with results from simulations based on a convection versus diffusion (C/D) model, with the endodermis being the main barrier for solutes and water. Solute profiles in the stele were calculated as they occurred during rapid water flows across the root. The model quantitatively predicted the experimental finding of two distinct phases during PRs, in terms of a build-up of concentration profiles in the stele between endodermis and xylem vessels. It also predicted that, following a PC, half-times (T1/2) of PRs increased as the time used for clamping (and the build-up of USLs) increased. Following PCs of durations of 15, 30 and 60 s, T1/2 increased by factors of between 2.5 and 7.0, and water permeability of roots (root hydraulic conductivity, Lpr) was reduced by the same factors. When root pressure was immediately taken back to the original equilibrium root pressure following a PC, there was a transient uptake of water into the root stele (transient increase of root pressure), and the size of transients rose with time of clamping, as predicted by the model. The results indicated that the 'real' hydraulic conductivity of roots should be measured during initial water flows, such as during the rapid phase of PRs, when the effect of USLs was minimized. It was discussed that 'pressure-propagation effects' could not explain the finding of two phases during PRs. The results of USL effects threw some doubt on the use of PC and high-pressure flowmeter (HPFM) techniques with roots, where rigorous estimates of USLs were still missing despite the fact that large quantities of water were forced across the root.

Plant size, not age, regulates growth and gas exchange in grafted Scots pine trees

We studied the effect of scion donor-tree age on the physiology and growth of 6- to 7-year-old grafted Scots pine (Pinus sylvestris L.) trees (4 and 5 years after grafting). Physiological measurements included photosynthethetic rate, stomatal conductance, transpiration, whole plant hydraulic conductance, needle nitrogen concentration and carbon isotope composition. Growth measurements included total and component biomasses, relative growth rates and growth efficiency. Scion donor trees ranged in age from 36 to 269 years at the time of grafting. Hydraulic conductance was measured gravimetrically, applying the Ohm's law analogy, and directly, with a high-pressure flow meter. We found no effect of scion donor-tree age on any of the variables measured. There was, however, great variation within scion donor-tree age groups, which was related to the size of the grafted trees. Differences in size may have been caused by variable initial grafting success, but there was no indication that grafting success and age were related. At the stem level, hydraulic conductance scaled with total leaf area so that total conductance per unit leaf area did not vary with crown size. However, leaf specific hydraulic conductance (gravimetric), transpiration, photosynthesis and stomatal conductance declined with increasing total tree leaf area and needle width. We hypothesize that needle width is inversely related to mesophyll conductance. We conclude that canopy and needle size and not scion donor-tree age determined gas exchange in our grafted trees.

Irradiance-induced plasticity in the hydraulic properties of saplings of different temperate broad-leaved forest tree species

We assessed the irradiance-related plasticity of hydraulic architecture in saplings of Betula pendula Roth., a pioneer species; Acer pseudoplatanus L., Fraxinus excelsior L. and Quercus robur L., which are post-pioneer light-requiring species; and Quercus petraea Matt. Liebl. and Fagus sylvatica L. Plants were grown in pots in 36%, 16% and 4% of full sunlight. Hydraulic conductance was measured with a high-pressure flow-meter in entire, in situ root systems and in excised shoots. Leaf-specific whole-plant conductance (LSC) increased with irradiance, due, in part, to an effect of irradiance on plant size. In addition, there was a size-independent effect of irradiance on LSC due, in part, to an increase in root hydraulic conductance paralleled by an increase in root biomass scaled to leaf area. Changes in shoot conductivity also contributed to the size-independent plasticity of LSC. Vulnerability to cavitation measured in current-year twigs was much larger in shade-grown plants. Betula pendula had the highest whole-plant, root and shoot conductances and also the greatest vulnerability to cavitation. The other species were similar in LSC, but showed some variation in root conductance scaled to biomass, with Q. robur, Q. petraea and F. sylvatica having the lowest root conductance and susceptibility to cavitation. All species showed a similar irradiance-related plasticity in LSC.

Effects of rootstock and irrigation regime on hydraulic architecture of &lt;em&gt;Vitis vinifera&lt;/em&gt; L. Cv. Tempranillo

&lt;p style="text-align: justify;"&gt;Rootstock hydraulic properties are probably the keys to drought or waterlogging resistance. This work evaluates the effects of two rootstocks (110R, SO4) and two irrigation levels (ET0, 0,5 ·ET0) on the hydraulic architecture of Tempranillo. The experiment was carried out on potted plants near Barcelona (Spain). Hydraulic resistance in trunks was measured by low pressure method and using a High Pressure Flow Meter (HPFM). The trunks of Tempranillo grafted onto 110R showed a lower resistance than the ones grafted onto SO4. The grafting point showed a higher resistance than the cultivar segment and the rootstock segment. HPFM results were more consistent than low-pressure measurements.&lt;/p&gt;

Circadian regulation of leaf hydraulic conductance in sunflower (Helianthus annuus L. cv Margot)

ABSTRACTThe circadian regulation of leaf hydraulic conductance (Kleaf) was investigated in Helianthus annuus L. (sunflower). Kleaf was measured with an high pressure flow meter during the light and dark period from plants growing at a photoperiod of 12 h. Kleaf was 4.0 e−4 kg s−1 m−2 MPa−1 during the light period (LL) and 30–40% less during the dark period (DL). When photoperiod was inverted and leaves were measured for Kleaf at their subjective light or dark periods, Kleaf adjusted to the new conditions requiring 48 h for increasing from dark to light values and 4 d for the opposite transition. Plants put in continuous dark showed Kleaf oscillating from light to dark values in phase with their subjective photoperiod indicating that Kleaf changes were induced by the circadian clock. Several cuts through the minor veins reduced leaf hydraulic resistance (Rleaf) of both LL and DL to the same value (1.0 e + 3 MPa m2 s kg−1) that equalled the vascular resistance (Rv). The contribution of the non‐vascular leaf resistance (Rnv) to Rleaf was of 71.9% in DL and of 58.4% in LL. The dominant Rnv was shown to be reversibly modulated by mercurials, suggesting that aquaporins play a role in diurnal changes of Kleaf.

The dependence of leaf hydraulic conductance on irradiance during HPFM measurements: any role for stomatal response?

This paper examines the dependence of whole leaf hydraulic conductance to liquid water (K(L)) on irradiance when measured with a high pressure flowmeter (HPFM). During HPFM measurements, water is perfused into leaves faster than it evaporates hence water infiltrates leaf air spaces and must pass through stomates in the liquid state. Since stomates open and close under high versus low irradiance, respectively, the possibility exists that K(L) might change with irradiance if stomates close tightly enough to restrict water movement. However, the dependence of K(L) on irradiance could be due to a direct effect of irradiance on the hydraulic properties of other tissues in the leaf. In the present study, K(L) increased with irradiance for 6 of the 11 species tested. Whole leaf conductance to water vapour, g(L), was used as a proxy for stomatal aperture and the time-course of changes in K(L) and g(L) was studied during the transition from low to high irradiance and from high to low irradiance. Experiments showed that in some species K(L) changes were not paralleled by g(L) changes. Measurements were also done after perfusion of leaves with ABA which inhibited the g(L) response to irradiance. These leaves showed the same K(L) response to irradiance as control leaves. These experimental results and theoretical calculations suggest that the irradiance dependence of K(L) is more consistent with an effect on extravascular (and/or vascular) tissues rather than stomatal aperture. Irradiance-mediated stimulation of aquaporins or hydrogel effects in leaf tracheids may be involved.

Hydraulic efficiency of the leaf venation system in sun- and shade-adapted species.

We tested the hypothesis that leaf hydraulics is correlated with the light adaptation of different plant species and specifically that the hydraulic resistance of the leaf venation (Rvenation) is lower in sun- than in shade-adapted species. Rvenation was measured in six sun- and six shade-adapted species with a high-pressure flow meter (HPFM). The number of conduits at the proximal third of the midrib was counted and the diameter of the widest conduits together with vein density were measured. Rvenation was higher in shade species than in sun species and it was negatively correlated with the mean diameter of the widest conduits. Maximum leaf conductance to water vapour recorded for the different species was negatively correlated with the corresponding Rvenation. Sun-adapted species coping with the high water demand typical of sunny habitats appeared to have developed a highly efficient conducting system to supply living mesophyll cells with water. In contrast, species adapted to shady habitats showed higher Rvenation values according to their lower need for investment of carbon into producing wide conduits in the leaf.

Bottlenecks to water transport in Quercus robur L.: the abscission zone and its physiological consequences

Abstract Cladoptosis, the abscission of twigs, is the main mechanism of changes in crown structure in senescing pedunculate oak (Quercus robur L.). We tested the hypotheses that abscission zones in nodes of old pedunculate oak trees reduce leaf-specific hydraulic conductance of shoots and thereby limit the stomatal conductance and assimilation. Hydraulic conductance and leaf-specific hydraulic conductance, measured with a high pressure flowmeter in 0.5–1.5 m long shoots, were significantly lower in shoots of low vigour compared to vigorous growing shoots in a 165-years-old stand in the southeast of Germany. Two types of bottlenecks to water transport could be identified in shoots of old oak trees, namely nodes and abscission zones. In young twigs, vessel diameter and vessel density in nodes with abscission zones were significantly reduced compared with internodes. In nodes without abscission zones, vessel density was significantly reduced. The reduction of hydraulic conductance was especially severe in the smallest and youngest shoots with diameters less than 2 mm. Internodes of 1–5 mm sapwood diameter had an average hydraulic conductance of 7.13×10 −6 ±0.2×10 −6 kg s −1 m −1 MPa−1, compared to 4.54×10 −6 ±0.3×10 −6 kg s −1 m −1 MPa−1 in those with nodes. Maximum stomatal conductance and maximum net assimilation rate increased significantly with hydraulic conductance and leaf-specific hydraulic conductance. Maximum rate of net photosynthesis Amax of the most vigorous shoots (VC0) ( 7.34±0.55 μmol m −2 s −1 ) was significantly higher (P 5.97±0.28 μmol m −2 s −1 ). Our data support the hypothesis that the changes in shoot and consequently crown architecture that are observed in ageing and declining trees can limit photosynthesis by reducing shoot hydraulic conductance. Abscission zones increase the hydraulic disadvantage of less vigorous compared to vigorously growing twigs. Cladoptosis might serve as a mechanism of selection between twigs of different efficiency.

Drought resistance of Ailanthus altissima: root hydraulics and water relations.

Drought resistance of Ailanthus altissima (Mill.) Swingle is a major factor underlying the impressively wide expansion of this species in Europe and North America. We studied the specific mechanism used by A. altissima to withstand drought by subjecting potted seedlings to four irrigation regimes. At the end of the 13-week treatment period, soil water potential was -0.05 MPa for well-watered control seedlings (W) and -0.4, -0.8 and -1.7 MPa for drought-stressed seedlings (S) in irrigation regimes S1, S2 and S3, respectively. Root and shoot biomass production did not differ significantly among the four groups. A progressively marked stomatal closure was observed in drought-stressed seedlings, leading to homeostasis of leaf water potential, which was maintained well above the turgor loss point. Root and shoot hydraulics were measured with a high-pressure flow meter. When scaled by leaf surface area, shoot hydraulic conductance did not differ among the treated seedlings, whereas root hydraulic conductance decreased by about 20% in S1 and S2 seedlings and by about 70% in S3 seedlings, with respect to the well-watered control value. Similar differences were observed when root hydraulic conductance was scaled by root surface area, suggesting that roots had become less permeable to water. Anatomical observations of root cross sections revealed that S3 seedlings had shrunken cortical cells and a multilayer endodermal-like tissue that probably impaired soil-to-root stele water transport. We conclude that A. altissima seedlings are able to withstand drought by employing a highly effective water-saving mechanism that involves reduced water loss by leaves and reduced root hydraulic conductance. This water-saving mechanism helps explain how A. altissima successfully competes with native vegetation.