The empirical calibration of Granier-type heat dissipation sap flow probes that relate temperature difference (DeltaT) to sap velocity (v) was reevaluated in stems of three tropical tree species. The original calibration was confirmed when the entire heated probe was in contact with conducting xylem, but mean v was underestimated when part of the probe was in contact with nonconducting xylem or bark. Analysis of the effects of nonuniform sap velocity profiles on heat dissipation estimates showed that errors increased as v and the proportion of the probe in nonconducting wood increased. If half of a 20-mm probe is in sapwood with a v of 0.15 mm s(-1) and the other half is in nonconducting wood, then mean v for the whole probe can be underestimated by as much as 50%. A correction was developed that can be used if the proportion of the probe in nonconducting wood is known. Even with the entire heated probe in contact with conducting xylem, v would be underestimated when radial velocity gradients are present. In this case, the error would be smaller except when velocity gradients are very steep, as can occur in species with ring-porous wood anatomy. Errors occur because the relationship between DeltaT and v is nonlinear. Mean DeltaT along the probe is therefore not a measure of mean v, and users of heat dissipation probes should not assume that v is integrated along the length of the probe. The same type of error can occur when DeltaT is averaged through time while v is changing, but the error is small unless there are sudden, step changes between zero and high sap velocity. It is recommended that relatively short probes (20 mm or less) be used and that probes longer than the depth of conducting sapwood be avoided. Multiple probes inserted to a range of depths should be used in situations where steep gradients in v are expected. If these conditions are met, heat dissipation probes remain useful and widely applicable for measuring sap flow in woody stems.
Heat and stable isotope tracers were used to study axial and radial water transport in relation to sapwood anatomical characteristics and internal water storage in four canopy tree species of a seasonally dry tropical forest in Panama. Anatomical characteristics of the wood and radial profiles of sap flow were measured at the base, upper trunk, and crown of a single individual of Anacardium excelsum, Ficus insipida, Schefflera morototoni, and Cordia alliodora during two consecutive dry seasons. Vessel lumen diameter and vessel density did not exhibit a consistent trend axially from the base of the stem to the base of the crown. However, lumen diameter decreased sharply from the base of the crown to the terminal branches. The ratio of vessel lumen area to sapwood cross-sectional area was consistently higher at the base of the crown than at the base of the trunk in A. excelsum, F. insipida and C. alliodora, but no axial trend was apparent in S. morototoni. Radial profiles of the preceding wood anatomical characteristics varied according to species and the height at which the wood samples were obtained. Radial profiles of sap flux density measured with thermal dissipation sensors of variable length near the base of the crown were highly correlated with radial profiles of specific hydraulic conductivity (k s ) calculated from xylem anatomical characteristics. The relationship between sap flux density and k s was speciesindependent. Deuterium oxide (D 2 O) injected into the base of the trunk of the four study trees was detected in the water transpired from the upper crown after only 1 day in the 26-m-tall C. alliodora tree, 2 days in the 28-m-tall F. insipida tree, 3 days in the 38-m-tall A. excelsum tree, and 5 days in the 22-m-tall S. morototoni tree. Radial transport of injected D 2 O was detected in A. excelsum, F. insipida and S. morototoni, but not C. alliodora. The rate of axial D 2 O transport, a surrogate for maximum sap velocity, was positively correlated with the predicted sapwood k s and with tree height normalized by the relative diurnal water storage capacity. Residence times for the disappearance of the D 2 O tracer in transpired water ranged from 2 days in C. alliodora to 22 days in A. excelsum and were positively correlated with a normalized index of diurnal water storage capacity. Capacitive exchange of water between stem storage compartments and the transpiration stream thus had a profound influence on apparent rates of axial water transport, the magnitude of radial water movement, and the retention time in the tree of water taken up by the roots. The inverse relationship between internal water exchange capacity and k s was consistent with a trade-off contributing to stability of leaf water status through highly efficient water transport at one extreme and release of stored water at the other extreme.
Robust thermal dissipation sensors of variable length (3 to 30 cm) were developed to overcome limitations to the measurement of radial profiles of sap flow in large-diameter tropical trees with deep sapwood. The effective measuring length of the custom-made sensors was reduced to 1 cm at the tip of a thermally nonconducting shaft, thereby minimizing the influence of nonuniform sap flux density profiles across the sapwood. Sap flow was measured at different depths and circumferential positions in the trunks of four trees at the Parque Natural Metropolitano canopy crane site, Panama City, Republic of Panama. Sap flow was detected to a depth of 24 cm in the trunks of a 1-m-diameter Anacardium excelsum (Bertero & Balb. ex Kunth) Skeels tree and a 0.65-m-diameter Ficus insipida Willd. tree, and to depths of 7 cm in a 0.34-m-diameter Cordia alliodora (Ruiz & Pav.) Cham. trunk, and 17 cm in a 0.47-m-diameter Schefflera morototoni (Aubl.) Maguire, Steyerm. & Frodin trunk. Sap flux density was maximal in the outermost 4 cm of sapwood and declined with increasing sapwood depth. Considerable variation in sap flux density profiles was observed both within and among the trees. In S. morototoni, radial variation in sap flux density was associated with radial variation in wood properties, particularly vessel lumen area and distribution. High variability in radial and circumferential sap flux density resulted in large errors when measurements of sap flow at a single depth, or a single radial profile, were used to estimate whole-plant water use. Diurnal water use ranged from 750 kg H2O day-1 for A. excelsum to 37 kg H2O day-1 for C. alliodora.
Despite the crucial role of carbon transport in whole plant physiology and its impact on plant-environment interactions and ecosystem function, relatively little research has tried to examine how phloem physiology impacts plant ecology. In this review, we highlight several areas of active research where inquiry into phloem physiology has increased our understanding of whole plant function and ecological processes. We consider how xylem-phloem interactions impact plant drought tolerance and reproduction, how phloem transport influences carbon allocation in trees and carbon cycling in ecosystems and how phloem function mediates plant relations with insects, pests, microbes and symbiotes. We argue that in spite of challenges that exist in studying phloem physiology, it is critical that we consider the role of this dynamic vascular system when examining the relationship between plants and their biotic and abiotic environment.
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