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Notizen: 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.

Notizen: 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.

Notizen: 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.

Notizen: It has been proposed that the stomatal response to humidity relies on sensing of the transpiration rate itself rather than relative humidity or the saturation deficit per se. We used independent measurements of stomatal conductance (gs), transpiration (E), and leaf-to-air vapour pressure difference (V) in a hybrid poplar canopy to evaluate relationships between gs and E and between gs and V. Relationships between E, V and total vapour phase conductance or crown conductance (gc) were also assessed. Conductance measurements were made on exposed and partially shaded branches over a wide range of incident solar radiation. In exposed branches, gs appeared to decline linearly with increasing E and increasing V at both high and low irradiance. However, in a partially shaded branch, a bimodal relationship between gs and E was observed in which gs continued to decrease after E had reached a maximum value and begun to decrease. The relationship between gs and V for this branch was linear. Plots of gc against E always yielded bimodal or somewhat variable relationships, whereas plots of gc against V were invariably linear. It was not possible to derive a unique relationship between conductance and E or V because prevailing radiation partially determined the operating range for conductance. Normalization of data by radiation served to linearize responses observed within the same day or type of day, but even after normalization, data collected on partly cloudy days could not be used to predict stomatal behaviour on clear days and vice versa. An additional unidentified factor was thus also involved in determining operating ranges of conductance on days with different overall radiation regimes. We suggest that the simplest mechanism to account for the observed humidity responses is stomatal sensing of the epidermal or cuticular transpiration rate rather than the bulk leaf or stomatal transpiration rate.

Notizen: Freezing resistance mechanisms were studied in five endemic Hawaiian species growing at high elevations on Haleakala volcano, Hawaii, where nocturnal subzero (°C) air temperatures frequently occur. Extracellular freezing occurred at around -5°C in leaves of Argyroxiphium sandwicense and Sophora chrysophylla, but these leaves can tolerate extracellular ice accumulation to -15°C and -12°C, respectively. Mucilage, which apparently acted as an ice nucleator, comprised 9 to 11% of the dry weight of leaf tissue in these two species. Leaves of Vaccinium reticulatum and Styphelia tameiameiae were also found to tolerate substantial extracellular freezing. Dubautia menziesii, on the other hand, exhibited the characteristics of permanent supercooling; a very rapid decline in liquid water content associated with simultaneous intracellular and extracellular freezing. However, in those species that tolerate extracellular freezing, the decline in liquid water content during freezing is relatively slow. Osmotic potential was lower at pre-dawn than at midday in four of the species studied. Nocturnal production of osmotically active solutes may have helped to prevent intracellular freeze dehydration as well as to provide non-colligative protection of cell membranes. Styphelia tameiameiae supercooled to -9·3°C and tolerated tissue freezing to below -15°C, a unique combination of physiological characteristics related to freezing. Tolerance of extracellular ice formation after considerable supercooling may have resulted from low tissue water content and a high degree of intracellular water binding in this species, as determined by nuclear magnetic resonance studies. The climate at high elevations in Hawaii is relatively unpredictable in terms of the duration of subzero temperatures and the lowest subzero temperature reached during the night. It appears that plants growing in this tropical alpine habitat have been under selective pressures for the evolution of freezing tolerance mechanisms.

Notizen: Stomatal regulation of transpiration was studied in hedgerow coffee (Coffea arabica L.) at different stages of canopy development encompassing a range of leaf area indices (L) from 0·7 to 6·7. Stomatal (gs) and crown (gc) conductance attained maximum values early during the day and then declined as both leaf-to-bulk air water vapour mole fraction difference (Va) and photosynthetically active photon flux density (I) continued to increase. Covariation of environmental variables during the day, particularly V, I, and wind speed (u), obscured stomatal responses to individual variables. This also caused diurnal hysteresis in the relationship between gc and individual variables. Normalization of gs and gc by I removed the hysteresis and revealed a strong stomatal response to humidity. At the crown scale, transpiration (E) increased linearly with net radiation (Rn) and seemed to increase with increasing wind speed. Increasing wind speed imposed higher leaf interior to leaf surface water vapour mole fraction differences (Vs) at given levels of Va. However, strong relationships between declining gc and E and increasing wind speed were obtained when gc and E were normalized by I and Rn, respectively, without invoking additional potential interactions involving temperature or CO2 concentration at the leaf surface. Apparent stomatal responses to wind were thus at least partially a reflection of the stomatal response to humidity.

Notizen: Since its introduction in the late 19th century, the so-called cohesion theory has become widely accepted as explaining the mechanism of the ascent of sap. According to the cohesion theory, the minimum standing vertical xylem tension gradient should be 0·01 MPa m−1. When transpiration is occurring, frictional resistances are expected to make this gradient considerably steeper. The results of numerous pressure chamber measurements reported in the literature are generally regarded as corroborating the cohesion theory. Nevertheless, several reports of pressure chamber measurements in tall trees appear to be incompatible with predictions of the cohesion theory. Furthermore, the pressure chamber is an indirect method for inferring xylem pressure, which, until recently, has not been validated by comparison against a direct method. The xylem pressure probe provides a means of testing the validity of the pressure chamber and other indirect techniques for estimating xylem pressure. We discuss here the results of concurrent measurements made with the pressure chamber and the xylem pressure probe, particularly recent measurements made at the top of a tall tropical tree during the rainy season. These measurements indicate that the pressure chamber often substantially overestimates the tension previously existing in the xylem, especially in the partially dehydrated tissue of droughted plants. We also discuss other evidence obtained from classical and recent approaches for studying water transport. We conclude that the available evidence derived from a wide range of independent approaches warrants a critical reappraisal of tension-driven water transport as the exclusive mechanism of long-distance water transport in plants.

Notizen: Hydraulic conductance was measured on leaf and stem segments excised from sugarcane plants at different stages of development. Maximum transpiration rates and leaf water potential (ΨL) associated with maximum transpiration were also measured in intact plants as a function of plant size. Leaf specific hydraulic conductivity (Lsc) and transpiration on a unit leaf area basis (E) were maximal in plants with approximately 0.2 m2 leaf area and decreased with increasing plant size. These changes in Fand Lsc were nearly parallel, which prevented φL in larger plants from decreasing to levels associated with substantial loss in xylem conductivity caused by embolism formation. Coordination of changes in E and leaf hydraulic properties was not mediated by declining leaf water status, since φL increased with plant size. Hydraulic constrictions were present at nodes and in the node-leaf sheath-leaf blade pathway. This pattern of constrictions is in accord with the idea of plant segmentation into regions differing in water transport efficiency and would tend to confine embolisms to the relatively expendable leaves at terminal positions in the pathway, thereby preserving water transport through the stem.

Notizen: 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.

Notizen: Stomatal control of crown transpiration was studied in Anacardium excelsum, a large-leaved, emergent canopy species common in the moist forests of Central and northern South America. A construction crane equipped with a gondola was used to gain access to the uppermost level in the crown of a 35-m-tall individual. Stomatal conductance at the single leaf scale, and transpiration and total vapour phase conductance (stomatal and boundary layer) at the branch scale were measured simultaneously using the independent techniques of porometry and stem heat balance, respectively. This permitted the sensitivity of transpiration to a marginal change in stomatal conductance to be evaluated using a dimensionless coupling coefficient (1-ω) ranging from zero to 1, with 1 representing maximal stomatal control of transpiration. Average stomatal conductance varied from 0.09 mol m−2 s−1 during the dry season to 0.3 mol m−2 s−1 during the wet season. Since boundary layer conductance was relatively low (0.4 mol m−2 s−1), 1-ω ranged from 0.46 during the dry season to only 0.25 during the wet season. A pronounced stomatal response to humidity was observed, which strongly limited transpiration as evaporative demand increased. The stomatal response to humidity was apparent only when the leaf surface was used as the reference point for measurement of external vapour pressure. Average transpiration was predicted to be nearly the same during the dry and wet seasons despite a 1 kPa difference in the prevailing leaf-to-air vapour pressure difference. The patterns of stomatal behaviour and transpiration observed were consistent with recent proposals that stomatal responses to humidity are based on sensing the transpiration rate itself.