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Notes: The pathway for water movement from the soil through plants to the atmosphere can be represented by a series of liquid and vapour phase resistances. Stomatal regulation of vapour phase resistance balances transpiration with the efficiency of water supply to the leaves, avoiding leaf desiccation at one extreme, and unnecessary restriction of carbon dioxide uptake at the other. In addition to maintaining a long-term balance between vapour and liquid phase water transport resistances in plants, stomata are exquisitely sensitive to short-term, dynamic perturbations of liquid water transport. In balancing vapour and liquid phase water transport, stomata do not seem to distinguish among potential sources of variation in the apparent efficiency of delivery of water per guard cell complex. Therefore, an apparent soil-to-leaf hydraulic conductance based on relationships between liquid water fluxes and driving forces in situ seems to be the most versatile for interpretation of stomatal regulatory behaviour that achieves relative homeostasis of leaf water status in intact plants. Components of dynamic variation in apparent hydraulic conductance in intact plants include, exchange of water between the transpiration stream and internal storage compartments via capacitive discharge and recharge, cavitation and its reversal, temperature-induced changes in the viscosity of water, direct effects of xylem sap composition on xylem hydraulic properties, and endogenous and environmentally induced variation in the activity of membrane water channels in the hydraulic pathway. Stomatal responses to humidity must also be considered in interpreting co-ordination of vapour and liquid phase water transport because homeostasis of bulk leaf water status can only be achieved through regulation of the actual transpirational flux. Results of studies conducted with multiple species point to considerable convergence with regard to co-ordination of stomatal and hydraulic properties. Because stomata apparently sense and respond to integrated and dynamic soil-to-leaf water transport properties, studies involving intact plants under both natural and controlled conditions are likely to yield the most useful new insights concerning stomatal co-ordination of transpiration with soil and plant hydraulic properties.

Notes: Abstract Leaf water relations characteristics of creosote bush, Larrea tridentata, were studied in view of previous reports that its leaves commonly experience zero or negative turgor under dry conditions. Leaf turgor loss point (〈inlineGraphic alt="inline image" href="urn:x-wiley:01407791:PCE467:PCE_467_mu1" location="equation/PCE_467_mu1.gif"/〉) was determined by a pressure-volume method for samples subjected to a hydration procedure and for untreated samples. Hydration caused 〈inlineGraphic alt="inline image" href="urn:x-wiley:01407791:PCE467:PCE_467_mu1" location="equation/PCE_467_mu1.gif"/〉 to increase by as much as 3 M Pa. Hydration of samples also caused changes in other leaf water relations characteristics such as symplastic solute content, tissue elasticity and symplasmic water fraction, but total leaf solute content was unchanged. Comparison of our field plant water potential data with values of 〈inlineGraphic alt="inline image" href="urn:x-wiley:01407791:PCE467:PCE_467_mu1" location="equation/PCE_467_mu1.gif"/〉 obtained by the two methods resulted in predictions of turgor loss during part or all of a diurnal cycle based on hydrated samples, and turgor maintenance (at least 0.3 MPa) based on untreated samples. Pooled data for 〈inlineGraphic alt="inline image" href="urn:x-wiley:01407791:PCE467:PCE_467_mu1" location="equation/PCE_467_mu1.gif"/〉 obtained from both partially hydrated and untreated samples showed that L. tridentata maintains fairly constant levels of turgor over a wide range of leaf water potential. Dilution of cell contents by apoplastic water introduced significant errors in psychrometric determinations of osmotic potential in both frozen and thawed leaf tissue and expressed cell sap. Use of these values of osmotic potential resulted in predictions of zero turgor at all plant water potentials measured in the field.

Notes: Abstract Stomatal conductance per unit leaf area in well-irrigated field- and greenhouse-grown sugarcane increased with leaf area up to 0.2 m2 plant 1, then declined so that maximum transpiration per plant tended to saturate rather than increase linearly with further increase in leaf area. Conductance to liquid water transport exhibited parallel changes with plant size. This coordiantion of vapour phase and liquid phase conductances resulted in a balance between water loss and water transport capacity, maintaining leaf water status remarkably constant over a wide range of plant size and growing conditions. The changes in stomatal conductance were not related to plant or leaf age. Partial defoliation caused rapid increases in stomatal conductance, to re-establish the original relationship with remaining leaf area. Similarly, pruning of roots caused rapid reductions in stomatal conductance, which maintained or improved leaf water status. These results suggest that sugarcane stomata adjusted to the ratio of total hydraulic conductance to total transpiring leaf area. This could be mediated by root metabolites in the transpiration stream, whose delivery per unit leaf area would be a function of the relative magnitudes of root system size, transpiration rate and leaf area.

Notes: Abstract. Gas exchange data obtained with wellventilated leaf cuvettes provide clear evidence of a stomatal response to leaf-air vapour pressure difference (V). In contrast, remotely sensed leaf temperatures with specific assumptions regarding canopy boundary layer characteristics, have been interpreted to mean that stomata do not respond to V. We address this apparent discrepancy in a sugarcane field by simultaneous application of a single-leaf, porometric technique and a whole-canopy, Bowen ratioenergy balance technique. These methods indicated significant stomatal response to V in well-irrigated sugarcane. Stomatal responses to V in the field were obscured by strong covariance of major environmental parameters so that opening responses to light and closing responses to V tended to offset each other. Low boundary layer conductance significantly uncoupled V at the leaf surface (Vs) from V determined in the bulk atmosphere (Va). This reduced the range of the stimulus, Vs, thereby reducing the range of the stomatal response, without indicating low stomatal sensitivity to V. Stomatal responses to Va may be smaller than expected from V response curves in cuvettes, since Vs rather than the conventionally measured Va is analogous to V in a well-stirred cuvette. Recently published conclusions that remotely sensed canopy temperatures are inconsistent with stomatal response to V may be based on erroneous estimates of canopy boundary layer conductance and thus of Vs, use of air saturation deficit rather than V to express evaporative demand, and investigation at higher levels of evaporative demand than those eliciting maximal stomatal response.

Notes: Diel variation in specific hydraulic conductivity (ks) was recorded in petioles of two savanna tree species, Schefflera macrocarpa and Caryocar brasiliense, from central Brazil. These two species have compound leaves with long petioles (10–30 cm). In both species, petiole ks decreased sharply with increasing transpiration rates and declining leaf water potentials (ψL) during the morning. Petiole ks increased during the afternoon while the plants were still transpiring and the water in the non-embolized vessels was still under tension. Dye experiments confirmed that in both species diel variation in ks was associated with embolism formation and repair. When transpiration was prevented in individual leaves, their petiole ks and water potential remained close to their maximum values during the day. When minimum daily ψL on selected branches was experimentally lowered by 0.2–0.6 MPa, the rate of ks recovery during the afternoon was slower in comparison with control branches. Several field manipulations were performed to identify potential mechanisms involved in the refilling of embolized petiole vessels. Removal of the cortex or longitudinal incisions in the cortex prevented afternoon recovery of ks and refilling of embolized vessels. When distilled water was added to petiole surfaces that had been abraded to partially remove the cuticle, ks increased sharply during the morning and early afternoon. Evidence of starch to sugar conversion in the starch sheath cells surrounding the vascular bundles of the petioles was observed during periods of rapid transpiration when the abundance of starch granules in the starch sheath cells surrounding the vascular bundles decreased. Consistent with this, petiole sugar content was highest in the early afternoon. The most parsimonious explanation of the field observations and the experimental results was that an increase in osmotically active solutes in cells outside the vascular bundles at around midday leads to water uptake by these cells. However, the concurrent increase in tissue volume is partially constrained by the cortex, resulting in a transient pressure imbalance that may drive radial water movement in the direction of the embolized vessels, thereby refilling them and restoring water flow. This study thus presents evidence that embolism formation and repair are two distinct phenomena controlled by different variables. The degree of embolism is a function of tension, and the rate of refilling a function of internal pressure imbalances.

Notes: Stem water storage capacity and diurnal patterns of water use were studied in five canopy trees of a seasonal tropical forest in Panama. Sap flow was measured simultaneously at the top and at the base of each tree using constant energy input thermal probes inserted in the sapwood. The daily stem storage capacity was calculated by comparing the diurnal patterns of basal and crown sap flow. The amount of water withdrawn from storage and subsequently replaced daily ranged from 4 kg d–1 in a 0·20-m-diameter individual of Cecropia longipes to 54 kg d–1 in a 1·02-m-diameter individual of Anacardium excelsum, representing 9–15% of the total daily water loss, respectively. Ficus insipida, Luehea seemannii and Spondias mombin had intermediate diurnal water storage capacities. Trees with greater storage capacity maintained maximum rates of transpiration for a substantially longer fraction of the day than trees with smaller water storage capacity. All five trees conformed to a common linear relationship between diurnal storage capacity and basal sapwood area, suggesting that this relationship was species-independent and size-specific for trees at the study site. According to this relationship there was an increment of 10 kg of diurnal water storage capacity for every 0·1 m2 increase in basal sapwood area. The diurnal withdrawal of water from, and refill of, internal stores was a dynamic process, tightly coupled to fluctuations in environmental conditions. The variations in basal and crown sap flow were more synchronized after 1100 h when internal reserves were mostly depleted. Stem water storage may partially compensate for increases in axial hydraulic resistance with tree size and thus play an important role in regulating the water status of leaves exposed to the large diurnal variations in evaporative demand that occur in the upper canopy of seasonal lowland tropical forests.

Notes: Concurrent, independent measurements of stomatal conductance (gs), transpiration (E) and microenvironmental variables were used to characterize control of crown transpiration in four tree species growing in a moist, lowland tropical forest. Access to the upper forest canopy was provided by a construction crane equipped with a gondola. Estimates of boundary layer conductance (gb) obtained with two independent methods permitted control of E to be partitioned quantitatively between gs and gb using a dimensionless decoupling coefficient (Ω) ranging from zero to 1. A combination of high gs (c. 300–600 mmol m−2 s−1) and low wind speed, and therefore relatively low gb (c. 100–800 mmol m−2 s−1), strongly decoupled E from control by stomata in all four species (Ω= 0.7–0.9). Photosynthetic water-use efficiency was predicted to increase rather than decrease with increasing gs because gb was relatively low and internal conductance to CO2 transfer was relatively high. Responses of gs to humidity were apparent only when the leaf surface, and not the bulk air, was used as the reference point for determination of external vapour pressure. However, independent measurements of crown conductance (gc), a total vapour phase conductance that included stomatal and boundary layer components, revealed a clear decline in gc with increasing leaf-to-bulk air vapour pressure difference (Va because the external reference points for determination of gc and Va were compatible. The relationships between gc and Vc and between gs and Vs appeared to be distinct for each species. However, when gs and gc were normalized by the branch-specific ratio of leaf area to sapwood area (LA/SA), a morphological index of potential transpirational demand relative to water transport capacity, a common relationship between conductance and evaporative demand for all four species emerged. Taken together, these results implied that, at a given combination of LA/SA and evaporative demand scaled to the appropriate reference point, the vapour phase conductance and therefore transpiration rates on a leaf area basis were identical in all four contrasting species studied.

Notes: Abstract. Stomatal conductance of single leaves and transpiration from an entire sugarcane (Saccharum spp. hybrid) canopy were measured simultaneously using independent techniques. Stomatal and environmental controls of transpiration were assessed at three stages of canopy development, corresponding to leaf area indices (L) of 2.2, 3.6 and 5.6. Leaf and canopy boundary layers impeded transport of transpired water vapour away from the canopy, causing humidity around the leaves to find its own value through local equilibration rather than a value determined by the humidity of the bulk air mass above the canopy. This tended to uncouple transpiration from direct stomatal control, so that transpiration predicted from measurement of stomatal conductance and leaf-to-air vapour pressure differences was increasingly overestimated as the reference point for ambient vapour pressure measurement was moved farther from the leaf and into the bulk air. The partitioning of control between net radiation and stomata was expressed as a dimensionless decoupling coefficent ranging from zero to 1.0. When the stomatal aperture was near its maximum this coefficient was approximately 0.9, indicating that small reductions in stomatal aperture would have had little effect on canopy transpiration. Maximum rates of transpiration were, however, limited by large adjustments in maximum stomatal conductance during canopy development. The product of maximum stomatal conductance and L. a potential total canopy conductance in the absence of boundary layer effects, remained constant as L increased. Similarly, maximum canopy conductance, derived from independent micrometeorological measurements, also remained constant over this period. Calculations indicated that combined leaf and canopy boundary layer conductance decreased with increasing L such that the ratio of boundary layer conductance to maximum stomatal conductance remained nearly constant at approximately 0.5. These observations indicated that stomata adjusted to maintain both transpiration and the degree of stomatal control of transpiration constant as canopy development proceeded.

Notes: The present study examines the manner in which several whole-tree water transport properties scale with species-specific variation in sapwood water storage capacity. The hypothesis that constraints on relationships between sapwood capacitance and other water relations characteristics lead to predictable scaling relationships between intrinsic capacitance and whole-tree behaviour was investigated. Samples of sapwood from four tropical forest canopy tree species selected to represent a range of wood density, tree size and architecture, and taxonomic diversity were used to generate moisture release curves in thermocouple psychrometer chambers, from which species-specific values of sapwood capacitance were calculated. Sapwood capacitance was then used to scale several whole-tree water transport properties determined from measurements of upper branch and basal sap flow, branch water potential, and axial and radial movement of deuterated water (D2O) injected into the base of the trunk as a tracer. Sapwood capacitance ranged from 83 to 416 kg m−3 MPa−1 among the four species studied and was strongly correlated with minimum branch water potential, soil-to-branch hydraulic conductance, daily utilization of stored water, and axial and radial movement of D2O. The species-independent scaling of several whole-tree water transport properties with sapwood capacitance indicated that substantial convergence in plant function at multiple levels of biological organization was revealed by a simple variable related to a biophysical property of water transport tissue.

Notes: Dwarf mistletoe (Arceuthobium spp.) is a hemiparasite that is said to be the single-most destructive pathogen of commercially valuable coniferous trees in many regions of the world. Although its destructive nature is well documented in many respects, its effects on the physiology of its host are poorly understood. In the present study, water and carbon relations were characterized over a range of scale from leaf to whole tree in large (40- to 50-m-tall) individuals of western hemlock (Tsuga heterophylla (Raf.) Sarg.) that were either heavily infected, or uninfected with hemlock dwarf mistletoe (Arceuthobium tsugense). Specific hydraulic conductivity (ks) of infected branches was approximately half that of uninfected branches, yet leaf-specific conductivity (kL) was similar because leaf area : sapwood area ratios (AL : AS) of infected branches were lower. Pre-dawn and minimum leaf water potential and stomatal conductance (gs) were similar among infected and uninfected trees because adjustments in hydraulic architecture of infected trees maintained kL despite reduced ks. Maximum whole-tree water use was substantially lower in infected trees (approximately 55 kg d−1) than in uninfected trees (approximately 90 kg d−1) because reduced numbers of live branches in infected trees reduced whole-tree AL : AS in a manner consistent with that observed in infected branches. Maximum photosynthetic rates of heavily infected trees were approximately half those of uninfected trees. Correspondingly, leaf nitrogen content was 35% lower in infected trees. Foliar δ13C values were 2.8‰ more negative in infected than in uninfected individuals, consistent with the absence of stomatal adjustment to diminished photosynthetic capacity. Adjustments in hydraulic architecture of infected trees thus contributed to homeostasis of water transport efficiency and transpiration on a leaf area basis, whereas both carbon accumulation and photosynthetic water use efficiency were sharply reduced at both the leaf and whole-tree scale.