High-pressure Flow Meter Research Articles

Combined effects of arbuscular mycorrhizas and light on water uptake of the neotropical understory shrubs, Piper and Psychotria.

• Root hydraulic conductance (Kr ) was measured for five understory shrub species of the neotropical moist forest to determine the effects of arbuscular mycorrhizas (AM) for both carbon-rich and carbon-limited host plants. • Kr was measured using a high pressure flow meter (HPFM) for potted plants grown in a factorial combination of AM fungi (presence/absence) and light (3.5 and 30% of full sun, low/high). • AM colonization improved Kr for the more shade-tolerant species plants when growing in low light. By contrast, water uptake efficiency of the light-demanding species was significantly decreased by AM fungi in high light. Regardless of AM colonization, light-demanding species had a lower capacity than shade-tolerant species to meet transpirational demands, and they allocated substantially more to fine root production relative to leaf area when colonized. • The differential effects of AM colonization and light on a species' root hydraulic conductance in relation to phylogeny and light adaptation demonstrate that AM fungi may be critical in determining early plant succession and community composition not only due to effects on nutrient uptake, but on water uptake as well.

Changes in leaf hydraulic conductance correlate with leaf vein embolism in Cercis siliquastrum L.

The impact of xylem cavitation and embolism on leaf (K leaf) and stem (K stem) hydraulic conductance was measured in current-year shoots of Cercis siliquastrum L. (Judas tree) using the vacuum chamber technique. K stem decreased at leaf water potentials (ΨL) lower than −1.0 MPa, while K leaf started to decrease only at ΨL <−1.5 MPa. Leaf infiltration under vacuum with Phloxine B revealed that minor veins underwent extensive embolism and became non-functional at ΨL <−1.5 MPa, thus indicating that leaf vein embolism was closely related to K leaf changes. Field measurements of leaf conductance to water vapour (g L) and ΨL showed that stomata closed when ΨL decreased below the ΨL threshold inducing loss of hydraulic conductance in the leaf. The partitioning of hydraulic resistances within shoots and leaves was measured using the high-pressure flow meter method. The ratio of leaf to shoot hydraulic resistance was about 0.8, suggesting that stem cavitation had a limited impact on whole shoot hydraulic conductance. We suggest that stomatal aperture may be regulated by the cavitation-induced reduction of hydraulic conductance of the soil-to-leaf water pathway which, in turn, strongly depends on the hydraulic architecture of the plant and, in particular, on leaf hydraulics.

Hydraulic conductivities of competing root systems of Grevillea robusta and maize in agroforestry

Models of water uptake in mixed stands of vegetation commonly assume that water is partitioned among competing root systems in proportion to relative root length densities. Such an approach assumes implicitly that roots of different species have equivalent hydraulic properties. This was tested for root systems of Grevillea robustaA. Cunn. and maize (Zea maysL.) at a semi-arid site in Kenya. The hydraulic conductances for roots of both species were measured in situat the scale of the whole root or root system using a high pressure flow meter (HPFM). Hydraulic conductivities (κr) were expressed per unit root length. Root lengths were estimated for maize plants by soil coring and for G. robustausing a fractal branching model calibrated against soil coring. Mean κr was 1.88×10−7 ±0.28×10−7kg s−1 MPa−1 m−1 for G. robustaand 1.25×10−7 ±0.13×10−7kg s−1 MPa−1 m−1 for maize. Values of κr were not significantly different (P<0.05), suggesting that the assumption of hydraulic equivalence for root systems of the two species may be valid, at least when hydrostatic gradients are the major driving force for water uptake. Differences in conductivities between these species could arise, however, because of variation in the hydraulic properties of roots not accounted for here, for example because of root age, phenology or responses to the soil environment.

Hydraulic conductance of peach trees grafted on rootstocks with differing size-controlling potentials

SummaryThe objective of this study was to evaluate the hydraulic conductance in various components of peach trees grown on rootstocks with contrasting size-controlling potentials. The experiment was conducted on ‘Crimson Lady’ peach trees grafted onto two rootstocks: a semi-dwarfing rootstock, K-146-44 (P. salicina × P. persica hybrid) and a vigorous rootstock, Nemaguard (seed-propagated Prunus persica × P. davidiana hybrid). Hydraulic conductance of leaves, stems, trunk, graft union, and root system was measured in the scion/rootstocks with a high-pressure flow meter. Total leaf area and dry-matter partitioning into main tree components were also determined. The above and below ground portions of trees grown on Nemaguard rootstock were 393% and 299% larger, respectively, than trees grown on K-146-44 rootstock. Whether trees were grafted on Nemaguard or K-146-44, the root system accounted for most of the resistance to water flow through the tree. When hydraulic conductance was expressed per unit leaf area, trees on K-146-44 rootstock had significantly lower root conductance than trees on Nemaguard rootstock. Rootstocks had no effect on hydraulic conductance through the scion or the graft union. Trees on K-146-44 also partitioned relatively more dry matter to roots and leaves and less to stems than trees on Nemaguard. The results suggest that high root resistance plays a central role in the dwarfing mechanism induced by the size-controlling rootstock.

Drought until death do us part: a case study of the desiccation-tolerance of a tropical moist forest seedling-tree, Licania platypus (Hemsl.) Fritsch.

Studies of the desiccation tolerance of 15-month-old Licania platypus (Hemsl.) Fritsch seedlings were performed on potted plants. Pots were watered to field capacity and then dehydrated for 23-46 d to reach various visible wilting stages from slightly-wilted to dead. Root hydraulic conductance, k(r), was measured with a high-pressure flow meter and whole-stem hydraulic conductance, k(ws), was measured by a vacuum chamber method. Leaf punches were harvested for measurement of leaf water potential by a thermocouple psychrometer and for measurement of fresh- and dry-weight. L. platypus was surprisingly desiccation-tolerant, suggesting that most species of central Panama may be well adapted to the seasonality of rainfall in the region. The slightly-wilted stage corresponded to leaf water potentials and relative water contents of -2.7 MPa and 0.85, respectively, but plants did not die until these values fell to -7.5 MPa and 0.14, respectively. As desiccation proceeded k(r) and k(ws) declined relative to irrigated controls, but k(ws) was more sensitive to desiccation than k(r). Values of k(ws) declined by 70-85% in slightly-wilted to dead plants, respectively. By comparison, k(r) showed no significant change in slightly-wilted plants and fell by about 50% in plants having severely-wilted to dead shoots.

Reduced photosynthesis in old oak (Quercus robur): the impact of crown and hydraulic architecture.

We tested the hypothesis that changes in crown architecture of old pedunculate oak trees (Quercus robur L. ssp. robur Kl. et Kr. et Rol.) reduce leaf specific hydraulic conductance of shoots, thereby limiting stomatal conductance and assimilation of affected shoots. At the end of summer 1999, hydraulic conductance and leaf specific hydraulic conductance, measured with a high-pressure flow meter in 0.5- to 1.5-m long shoots, were 27 and 39% lower, respectively, in shoots of low vigor compared with vigorously growing shoots in a 165-year- old stand in southeastern Germany. Two types of bottlenecks to water transport can be identified in shoots of old oak trees, namely nodes and abscission zones. The reduction in hydraulic conductance was especially severe in shoots with diameters of less than 2 mm. Maximum stomatal conductance and maximum net assimilation rate increased significantly with hydraulic conductance and leaf specific hydraulic conductance. Our data support the hypothesis that changes in shoot and consequently crown architecture observed in aging trees can limit photosynthesis by reducing shoot hydraulic conductance. Thus, in addition to increasing pathway length and lower conductivity of xylem in old trees, structural changes in shoot and crown architecture need to be considered when analyzing water relations and photosynthesis in mature and declining trees.

Hydraulic conductance of root and shoot measured with the transient and dynamic modes of the high-pressure flowmeter

The hydraulic conductance (k) of shoots and root systems was measured using the transient and the dynamic modes of the high pressure flowmeter (HPFM). Measurements were conducted on Quercus robur and Fagus sylvatica plants grown on different subs- trates (forest soil, sand, Terra-green and vermiculite) and harvested at different times of the year. The values of k obtained by the tran- sient mode were compared to those obtained by the dynamic mode. A tight 1:1 correlation was observed for shoots and defoliated stems but several types of discrepancies appeared for root systems. The underestimation of k by the dynamic mode as compared to the transient mode could be explained by reverse osmosis at the endodermis. However the transient mode was not functional for some root systems. This problem occurred essentially in small plants harvested early in the year before budbreak had been completed. Nature and origins of problems are discussed. hydraulic conductance / high pressure flowmeter / transient mode / root / shoot

Relationship between hydraulic resistance and leaf morphology in broadleaf Quercus species: a new interpretation of leaf lobation

We investigated the relationship between leaf shape and leaf hydraulic resistance in a set of broadleaf Quercus tree species (Q. cerris, Q. frainetto, Q. petraea, Q. pyrenaica, Q. robur, Q. rubra, Q. velutina). Seedlings of all the studied species were grown under uniform environmental conditions. A new high-pressure flowmeter was designed to measure leaf-blade hydraulic resistance. Leaf shape was characterised by the complexity of leaf outline which was regarded as an estimate of leaf lobation. This was done using the box-counting fractal dimension of the leaf silhouette. Leaf hydraulic resistance was negatively related to leaf lobation. It is suggested that the lower hydraulic resistance in deeply lobed leaves may constitute a mechanism for improving water balance under dry atmospheric conditions.

A new high-pressure flowmeter for the estimation of hydraulic resistance in whole shoots of woody plants

A new high-pressure flowmeter has been developed and used to estimate hydraulic resistance in whole shoots of woody plants. It is based on the perfusion of pressurized water in the plant, and the simultaneous measurement of water flow (introduced volume of water per unit of time). This device was first calibrated, and then used to estimate the hydraulic resistance of stems and leaves in 1-yr seedlings of broadleaf Quercus species (Q. rubra, Q. cerris, Q. velutina, Q. petraea, Q. frainetto, Q. pyrenaica). The measured values are reliable and similar to those reported in the literature, but taking into account that this study dealt with seedlings. We have found significant differences between species and groups of species according to their hydraulic resistance values for stems and leaves. Those species whose seedlings showed lower leaf hydraulic resistance values can rehydrate their leaves more quickly and survive in environments with certain water stress (e.g., Q. pyrenaica). The high leaf hydraulic resistance of other species indicate that they can not resist these levels of water stress (e.g, Q. rubra). This study provided new insight to the importance of leaf hydraulic resistance as an adaptative trait to face water stress. Hydraulic resistance variability must be studied in depth as an additional mechanism within a wide array of morphological and ecophysiological adaptations that provide tree species a greater drought tolerance.

Xylem cavitation in the leaf of Prunus laurocerasus and its impact on leaf hydraulics.

This paper reports how water stress correlates with changes in hydraulic conductivity of stems, leaf midrib, and whole leaves of Prunus laurocerasus. Water stress caused cavitation-induced dysfunction in vessels of P. laurocerasus. Cavitation was detected acoustically by counts of ultrasonic acoustic emissions and by the loss of hydraulic conductivity measured by a vacuum chamber method. Stems and midribs were approximately equally vulnerable to cavitations. Although midribs suffered a 70% loss of hydraulic conductance at leaf water potentials of -1.5 MPa, there was less than a 10% loss of hydraulic conductance in whole leaves. Cutting and sealing the midrib 20 mm from the leaf base caused only a 30% loss of conduction of the whole leaf. A high-pressure flow meter was used to measure conductance of whole leaves and as the leaf was progressively cut back from tip to base. These data were fitted to a model of hydraulic conductance of leaves that explained the above results, i.e. redundancy in hydraulic pathways whereby water can flow around embolized regions in the leaf, makes whole leaves relatively insensitive to significant changes in conductance of the midrib. The onset of cavitation events in P. laurocerasus leaves correlated with the onset of stomatal closure as found recently in studies of other species in our laboratory.

Water relations of coastal and estuarine Rhizophora mangle: xylem pressure potential and dynamics of embolism formation and repair.

Physiological traits related to water transport were studied in Rhizophora mangle (red mangrove) growing in coastal and estuarine sites in Hawaii. The magnitude of xylem pressure potential (P x), the vulnerability of xylem to cavitation, the frequency of embolized vessels in situ, and the capacity of R. mangle to repair embolized vessels were evaluated with conventional and recently developed techniques. The osmotic potential of the interstitial soil water (πsw) surrounding the roots of R. mangle was c. -2.6±5.52×10-3 and -0.4±6.13×10-3MPa in the coastal and estuarine sites, respectively. Midday covered (non-transpiring) leaf water potentials (ΨL) determined with a pressure chamber were 0.6-0.8MPa more positive than those of exposed, freely-transpiring leaves, and osmotic potential of the xylem sap (πx) ranged from -0.1 to -0.3MPa. Consequently, estimated midday values of P x (calculated by subtracting πx from covered ΨL) were about 1MPa more positive than ΨL determined on freely transpiring leaves. The differences in ΨL between covered and transpiring leaves were linearly related to the transpiration rates. The slope of this relationship was steeper for the coastal site, suggesting that the hydraulic resistance was larger in leaves of coastal R. mangle plants. This was confirmed by both hydraulic conductivity measurements on stem segments and high-pressure flowmeter studies made on excised leafy twigs. Based on two independent criteria, loss of hydraulic conductivity and proportions of gas- and liquid-filled vessels in cryo-scanning electron microscope (cryo-SEM) images, the xylem of R. mangle plants growing at the estuarine site was found to be more vulnerable to cavitation than that of plants growing at the coastal site. However, the cryo-SEM analyses suggested that cavitation occurred more readily in intact plants than in excised branches that were air-dried in the laboratory. Cryo-SEM analyses also revealed that, in both sites, the proportion of gas-filled vessels was 20-30% greater at midday than at dawn or during the late afternoon. Refilling of cavitated vessels thus occurred during the late afternoon when considerable tension was present in neighboring vessels. These results and results from pressure-volume relationships suggest that R. mangle adjusts hydraulic properties of the water-transport system, as well as the leaf osmotic potential, in concert with the environmental growing conditions.

Limitation of stomatal conductance by hydraulic traits: sensing or preventing xylem cavitation?

We tested the hypothesis that hydraulic conductance per unit leaf surface area of plant shoots (KSL) determines the maximum diurnal stomatal conductance (gL) that can be reached by plants growing in the field. A second hypothesis was tested that some xylem cavitation cannot be avoided by transpiring plants and might act as a signal for regulating gL. Eleven woody species were studied, differing from each other with respect to taxonomy, wood anatomy and leaf habit. Maximum diurnal gL, transpiration rate (EL), pre-dawn and minimum diurnal leaf water potential (Ψpd and Ψmin, respectively) were measured in the field. The critical Ψ level at which stem cavitation was triggered (Ψcav) was measured on detached branches, using the acoustic method. A high-pressure flow meter was used to measure maximum KSL of 1-year-old shoots. Both gL and EL were positively related to KSL. The whole-plant hydraulic conductance per unit leaf area (KWL) of all the species studied, calculated as the ratio of EL to ΔΨ (=Ψpd-Ψmin) was closely related to KSL. In every case, Ψmin (ranging between –0.85 and –1.35 MPa in the different species) dropped to the Ψcav range or was <Ψcav (ranging between –0.71 and –1.23 MPa), thus suggesting that some cavitation-induced embolism could not be avoided. The possibility is discussed that some cavitation-induced reduction in KSL is the signal for stomatal closure preventing runaway embolism. The lack of correlation of gL to Ψcav is discussed in terms of the inconsistency of Ψcav as an indicator of the vulnerability of plants to cavitation. No differences in hydraulic traits were observed between evergreen and deciduous species.

Temperature effects on hydraulic conductance and water relations of Quercus robur L.

The effects of temperature on root and shoot hydraulic conductances (g(shoot) and g(root)) were investigated for Quercus robur L. saplings. In a first experiment, conductances were measured with a High Pressure Flow Meter on excised shoots and detopped root systems. The g(root) and g(shoot) increased considerably with temperature from 0-50 degrees C. Between 15 degrees C and 35 degrees C, g(shoot) and g(root) varied with water viscosity. In a second experiment, the impact of temperature-induced changes in g(root) on sapling transpiration (E) and leaf water potential (psileaf) was assessed. Intact plants were placed in a growth cabinet with constant air and variable soil temperatures. E increased linearly with soil temperature but psileaf remained constant. As a consequence, a linear relationship was found between E and g(plant). The results illustrate the significance of g(plant) for the stomatal control of transpiration and the significance of temperature for tree water transport.

Plant hydraulic conductance measured by the high pressure flow meter in crop plants

A new high pressure flow meter (HPFM) method for measuring plant hydraulic conductances (K) was investigated to examine whether its results are comparable to those from a conventional evaporative flux (EF) method in crops. Hydraulic conductance (K) was measured by the two methods under quasi-steady-state conditions in six crops grown in pots: soybean (Glycine max L. Merr. cv. Tsurunoko daizu), sunflower (Helianthus annuus L. cv. Russian mammoth), kidney bean (Phaseolus vulgaris L. cv. Tsurunashi morocco), tomato (Lycopersicon esculentum Mill. cv. Sekai-ichi), green pepper (Capsicum annuum L. cv. shishitou), and eggplant (Solanum melongena L. cv. Seiguro chunaga nasu). There was a 1:1 agreement between K values measured by the two methods for K values of whole plant, root and stem, and leaf under quasi-steady-state conditions. Leaf water potential (psi leaf) and evaporative flux density (E) in sunflower was curvilinear, indicating whole plant K estimated by the EF method increased with increase of E. Predicted psi leaf (= E divided by whole plant K measured by the HPFM method) agreed with measured psi leaf. Diurnal changes were also found in K measured by the HPFM confirming that K changed in response to temperature and E. The HPFM revealed that variable conductance was located in all organs: roots, stems, petioles, and leaves. These observations indicated that the HPFM is valid for crops as well as for trees (as previously established by Tsuda and Tyree) and has advantages over the EF method because of the speed and ease of the HPFM method.

Plant hydraulic conductance measured by the high pressure flow meter in crop plants

Plant hydraulic conductance measured by the high pressure flow meter in crop plants

Direct measurement of xylem pressure in leaves of intact maize plants. A test of the cohesion-tension theory taking hydraulic architecture into consideration

The water relations of maize (Zea mays L. cv Helix) were documented in terms of hydraulic architecture and xylem pressure. A high-pressure flowmeter was used to characterize the hydraulic resistances of the root, stalk, and leaves. Xylem pressure measurements were made with a Scholander-Hammel pressure bomb and with a cell pressure probe. Evaporation rates were measured by gas exchange and by gravimetric measurements. Xylem pressure was altered by changing the light intensity, by controlling irrigation, or by gas pressure applied to the soil mass (using a root pressure bomb). Xylem pressure measured by the cell pressure probe and by the pressure bomb agreed over the entire measured range of 0 to -0.7 MPa. Experiments were consistent with the cohesion-tension theory. Xylem pressure changed rapidly and reversibly with changes in light intensity and root-bomb pressure. Increasing the root-bomb pressure increased the evaporation rate slightly when xylem pressure was negative and increased water flow rate through the shoots dramatically when xylem pressure was positive and guttation was observed. The hydraulic architecture model could predict all observed changes in water flow rate and xylem. We measured the cavitation threshold for oil- and water-filled pressure probes and provide some suggestions for improvement.

Hydraulic architecture and water use of selected species from a lower montane forest in Panama

Plant water relations of nine woody species were studied in a lower montane rain forest in Panama. These data provide a partial test of the hypothesis that hydraulic architecture of lower montane species might limit transpiration and thus leaf size or nutrient transport (as suggested by J. Cavelier and E. G. Leigh, respectively). Diurnal variation in leaf transpiration was closely correlated with changes in net radiation. Peak transpiration rates (7 × 10–5 kg s–1 m–2) were as high as peak transpiration rates from tropical lowland forests but mean daily water use [0.39 ± 0.08 (SEM) kg m–2 day–1] were mostly lower than comparable data from tropical lowland forests. Thus transpiration rates are sufficiently high for sufficiently long periods to make it unlikely that nutrient transport is limited by transpiration. Another objective of this study was a comparison of two different methods to measure hydraulic conductance (Kh = flow rate per unit pressure gradient) and leaf specific conductance of stem segments (KL = Kh/leaf area distal to the segment). The results obtained with the traditional conductivity apparatus and the high pressure flow meter method, yielded similar results in six out of seven cases.

Whole-plant hydraulic resistance and vulnerability segmentation in Acer saccharinum.

Hydraulic properties were studied in Acer saccharinum L., a riparian species that also grows well on a dry soil when transplanted. Hydraulic resistances were measured by two independent techniques: a new high-pressure flowmeter (HPFM) method and a conventional evaporative flux (EF) method. Vulnerability to cavitation was also investigated on petioles, stems and roots using a hydraulic conductivity technique. Vulnerability segmentation was found, i.e., roots, stems and petioles had different vulnerabilities to xylem dysfunction. Petioles were most vulnerable with 50% loss of hydraulic conductivity at -0.5 MPa, roots were least vulnerable (50% loss at -2.2 MPa) and stems were intermediate in vulnerability. The HPFM and the EF methods gave comparable results, except that the EF method gave a significantly higher value for resistance across petioles plus leaves. Native embolism was high enough to explain the discrepancy in resistance across petioles plus leaves between the HPFM and the EF methods, indicating that the HPFM estimates the minimum (potential) hydraulic resistance of plants. Whole-plant hydraulic resistance of A. saccharinum was low compared to resistances of other temperate species. The hydraulic characteristics of A. saccharinum were consistent with adaptation to its typical environment: low whole-plant resistance assures high transpiration rates in the presence of sufficient water, and vulnerability segmentation provides the ability to survive during droughts through shedding of expendable organs.

Developmental control of xylem hydraulic resistances and vulnerability to embolism in Fraxinus excelsior L.: impacts on water relations

The hydraulic properties and leaf gas exchanges of Fraxinus excelsior L. branches differing by their age and their vertical crown position, but in comparable ambient air conditions (vapour pressure deficit and global radiation) were compared. The variations in leaflet water potential ψleaflet, leaflet stomatal conductance and transpiration rate, E, were small between different branches of the same crown. Whole branch hydraulic resistances (r branch ), and partitioning between leaf (r leaf ) and xylem resistance (r xylem ) were assessed with a high pressure flowmeter. r leaf represented 90% and 10% of r branch for upper and lower crown branches, respectively. The changes resulted from increases in r xylem caused by the formation of short shoot internodes mostly located in secondary axes. However, leaf area-specific branch resistances (r branch =r branch x LA) were nearly constant throughout the crown. This was consistent with the vertical variations in ψ leaflet because r branch x E represents the water potential drop from the trunk to the leaves. Because r xylem was higher, lower ψxylem values were predicted in lower crown rachises. However, rachises from lower crown branches were less vulnerable to embolism than in upper branches ( ψxylem at onset of embolism, ψ cav, were -3 and -2MPa, respectively). It was concluded that r xylem increased with branch age, but r* branch remained constant because LA decreased. As a consequence, E was maximized and ψxylem remained above ψcav, This suggested that, in Fraxinus, leaf gas exchanges and leaf areas were coupled with xylem hydraulic capacities probably through a control of bud activity.

Dynamic measurements of roots hydraulic conductance using a high-pressure flowmeter in the laboratory and field

A new high-pressure flowmeter (HPFM) is described which is capable of rapid water-flow measurements. The HPFM permits dynamic determination of hydraulic conductance of roots, K r , and can be used in the laboratory or field. The base of a root is connected to the HPFM and water is perfused into the root system opposite to the normal direction of flow during transpiration. The perfusion pressure is changed at a constant rate of 3-7 kPa s −1 while measuring the flow into the root every 2-4 s. The slope of the plot of flow versus applied pressure is K r . This paper describes the HPFM, presents the theory of dynamic flow measurements, discusses sources of error, presents evidence that dynamic measurements of K r in Ficus maclellandi (and six other tropical species from Panama) yield the correct result, and demonstrates the use of the method under field conditions in Panama on Cecropia obtusifolia and Palicourea guianensis