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1.
A dynamic model for simulating water flow in a Scots pine (Pinus sylvestris L.) tree was developed. The model is based on the cohesion theory and the assumption that fluctuating water tension driven by transpiration, together with the elasticity of wood tissue, causes variations in the diameter of a tree stem and branches. The change in xylem diameter can be linked to water tension in accordance with Hookea s law. The model was tested against field measurements of the diurnal xylem diameter change at different heights in a 37-year-old Scots pine at Hyyti?l?, southern Finland (61 degrees 51' N, 24 degrees 17' E, 181 m a.s.l.). Shoot transpiration and soil water potential were input data for the model. The biomechanical and hydraulic properties of wood and fine root hydraulic conductance were estimated from simulated and measured stem diameter changes during the course of 1 day. The estimated parameters attained values similar to literature values. The ratios of estimated parameters to literature values ranged from 0.5 to 0.9. The model predictions (stem diameters at several heights) were in close agreement with the measurements for a period of 6 days. The time lag between changes in transpiration rate and in sap flow rate at the base of the tree was about half an hour. The analysis showed that 40% of the resistance between the soil and the top of the tree was located in the rhizosphere. Modeling the water tension gradient and consequent woody diameter changes offer a convenient means of studying the link between wood hydraulic conductivity and control of transpiration.  相似文献   

2.
We examined interrelated xylem water tensions and embolism dynamics under field conditions by simultaneously monitoring ultra-acoustic emissions and changes in stem xylem diameter. Variation in stem xylem diameter was measured with linear displacement transducers to estimate variation in sap tension. Measured ultrasonic acoustic emissions coincided well with changes in xylem diameter, indicating that individual peaks in embolism occurred simultaneously with peaks in water tension. The good time resolution between measurements makes this method especially suitable for observing embolism dynamics on a short timescale. Longer lasting measurements can also be made to monitor inter-daily patterns in water tension and embolism because the techniques are non-destructive. Ultra-acoustic emissions occurred mainly during periods of decreasing stem xylem diameter, i.e., increasing water tension, when the water tension was high enough. Embolism also occurred during periods of increasing xylem diameter, i.e., decreasing water tension, but the number of embolizing conduits under these conditions was small.  相似文献   

3.
Alméras T 《Tree physiology》2008,28(10):1513-1523
Tree stems shrink in diameter during the day and swell during the night in response to changes in water tension in the xylem. Stem shrinkage can easily be measured in a nondestructive way, to derive continuous information about tree water status. The relationship between the strain and the change in water tension can be evaluated by empirical calibrations, or can be related to the structure of the plant. A mechanical analysis was performed to make this relationship explicit. The stem is modeled as a cylinder made of multiple layers of tissues, including heartwood, sapwood, and inner and outer bark. The effect of changes in water tension on the apparent strain at the surface of a tissue is quantified as a function of parameters defining stem anatomy and the mechanical properties of the tissues. Various possible applications in the context of tree physiology are suggested.  相似文献   

4.
In a mature beech stand located in north-eastern Germany, xylem sap flux measurements were continuously performed during the 2002–2004 growing seasons. Ten representative trunks were studied using heated thermal dissipation probes. The measurements aimed at identifying principles governing radial profiles of xylem flux in beech trunks. The measurements were taken up to a trunk depth of 132 mm. The sap flow density in the pericambial xylem was found to vary among trees of different diameters, but was not considerably smaller in suppressed trees. A model for the radial distribution of sap flux density was formulated relating trunk radius and sap flow density. The model takes into account different trunk diameter. About 90% of the sap flux was found to occur in the outer two fifths of the trunk. Using this model, an adequate estimate of transpiration can be achieved at tree and stand level, even when the sap flux measurements are restricted to the outer trunk sectors.  相似文献   

5.
Variations in radial patterns of xylem water content and sap flow rate were measured in five laurel forest tree species (Laurus azorica (Seub.) Franco, Persea indica (L.) Spreng., Myrica faya Ait., Erica arborea L. and Ilex perado Ait. ssp. platyphylla (Webb & Berth.) Tutin) growing in an experimental plot at Agua García, Tenerife, Canary Islands. Measurements were performed around midday during warm and sunny days by the heat field deformation method. In all species, water content was almost constant (around 35% by volume) over the whole xylem cross-sectional area. There were no differences in wood color over the whole cross-sectional area of the stem in most species with the exception of E. arborea, whose wood became darker in the inner layers. Radial patterns of sap flow were highly variable and did not show clear relationships with tree diameter or species. Sap flow occurred over the whole xylem cross-sectional area in some species, whereas it was limited to the outer xylem layers in others. Sap flow rate was either similar along the xylem radius or exhibited a peak in the outer part of the xylem area. Low sap flow rates with little variation in radial pattern were typical for shaded suppressed trees, whereas dominant trees exhibited high sap flow rates with a peak in the radial pattern. Stem damage resulted in a significant decrease in sap flow rate in the outer xylem layers. The outer xylem is more important for whole tree water supply than the inner xylem because of its larger size. We conclude that measurement of radial flow pattern provides a reliable method of integrating sap flow from individual measuring points to the whole tree.  相似文献   

6.
In plant physiology models involving bubble nucleation, expansion or elimination, it is typically assumed that the surface tension of xylem sap is equal to that of pure water, though this has never been tested. In this study we collected xylem sap from branches of the tree species Populus tremuloides, Betula papyrifera and Sorbus aucuparia over 3 months. We measured the instantaneous surface tension and followed changes over a period of 0.5-5 h using the pendant drop technique. In all three species the instantaneous surface tension was equal to or within a few percent of that of pure water. Further, in B. papyrifera and S. aucuparia the change over time following drop establishment, although significant, was very small. In P. tremuloides, however, there was a steep decline in surface tension over time that leveled off towards values 21-27% lower than that of pure water. This indicated the presence of surfactants. The values were lower for thinner distal branch segments than for proximal ones closer to the trunk. In some species it appears valid to assume that the surface tension of xylem sap is equal to that of water. However, in branch segments of P. tremuloides close to the terminal bud and hence potentially in other species as well, it may be necessary to take into account the presence of surfactants that reduce the surface tension over time.  相似文献   

7.
We investigated the vulnerability of xylem to embolism and the seasonal occurrence of xylem embolism in Italian alder (Alnus cordata Loisel.) by acoustic and hydraulic methods. Wood anatomy was also studied. More than eighty percent of the vessels were less than 50 mm long and no vessels were longer than 120 mm. Mean vessel diameter was 48 micro m. Ultrasound acoustic emissions from root and branch segments dehydrating in air followed a similar pattern: in both tissues, emission peaks were recorded when the relative water content of the xylem was around 0.2. In branches dehydrating in air, xylem embolism increased linearly as water potential decreased. In trees in the field, more than 80 percent of hydraulic conductivity was lost in the tree crowns during winter. Recovery from winter embolism occurred mostly before bud burst. In summer, xylem embolism was low (< 30%) and acoustic emissions from roots, stem and branches of trees in the field were also low.  相似文献   

8.
Mountain ash (Eucalyptus regnans F.J. Muell.) forest catchments exhibit a strong relationship between stand age and runoff, attributed inter alia to differences in tree water use. However, the tree water use component of the mountain ash forest water balance is poorly quantified. We have used the sap flow technique to obtain estimates of daily water use in large mountain ash trees. First, the sap flow technique was validated by means of an in situ cut tree experiment. Close agreement was obtained between the sap flow estimate of water use and the actual uptake of water by the tree from a reservoir. Second, we compared the variability in sap velocity between a symmetric and an asymmetric tree by using multiple sap flow loggers. In the symmetric tree, velocity was fairly uniform throughout the xylem during the day, indicating that accurate sap flow estimates can be obtained with a minimal number of sampling points. However, large variations in sap velocity were observed in the asymmetric tree, indicating that much larger sampling sizes are required in asymmetric stems for an accurate determination of mean sap velocity. Finally, we compared two procedures for scaling individual tree sap flow estimates to the stand level based on stem diameter and leaf area index measurements. The first procedure was based on a regression between stem diameter and tree water use, developed on a small sample of trees and applied to a stand-level census of stem diameter values. Inputs to the second procedure were tree water use and leaf area of a single tree and the leaf area index of the stand. The two procedures yielded similar results; however, the first procedure was more robust but it required more sampling effort than the second procedure.  相似文献   

9.
Trees of tropical semi-deciduous forests range from "drought-avoiding" stem-succulent species with low-density wood (< 0.5 g cm(-3)), which maintain high stem water potentials (psi(STEM) > -0.7 MPa) throughout the year, to "drought-tolerant" deciduous hardwood species (wood density > 0.75 g cm(-3)), which dehydrate strongly during seasonal drought (psi(STEM) < -6 MPa). In stem-succulent and other drought-avoiding species, xylem vessels are surrounded by extensive parenchyma providing intracellular water storage, whereas in deciduous species stem water storage is mainly extracellular. Thirteen tropical and two temperate tree species, representing different functional types, were studied. The contribution of stem water storage to these species' water use during water stress was determined by time-series analysis of dehydration and rehydration of excised leaf-bearing branches of these trees. During dehydration, stem water potential slowly declined 1-2 MPa in drought-avoiding species, but in deciduous species it rapidly fell 4-5 MPa, suggesting that water storage capacitance was related to xylem anatomy. After immersion of dehydrated, leafless branches in water, the decline in xylem tension and rate of water uptake during rehydration were linearly related, as predicted by application of Ohm's law to water flux. The decline of xylem tension during rehydration was biphasic, with a phase of rapid water uptake into extracellular spaces being followed by a prolonged phase of slow water uptake into living cells. The rate of water uptake during rehydration and the minima of leaf water potential observed in the field during the dry season were highly correlated with water storage capacitance, indicating that wood anatomy is a major determinant of drought adaptation.  相似文献   

10.
The freezing pattern and frost killing temperatures of apple (Malus domestica Borkh.) xylem were determined by differential thermal analysis and infrared differential thermal analysis (IDTA). Results from detached or attached twigs in controlled freezing experiments and during natural field freezing of trees were compared. Non-lethal freezing of apoplastic water in apple xylem as monitored during natural winter frosts in the field occurred at -1.9?±?0.4 °C and did not change seasonally. The pattern of whole tree freezing was variable and specific to the environmental conditions. On detached twigs high-temperature freezing exotherms (HTEs) occurred 2.8 K below the temperature observed under natural frosts in the field with a seasonal mean of -4.7?±?0.5 °C. Microporous apple xylem showed freezing without a specific pattern within a few seconds in IDTA images during HTEs, which is in contrast to macroporous xylem where a 2D freezing pattern mirrors anatomical structures. The pith tissue always remained unfrozen. Increasing twig length increased ice nucleation temperature; for increased twig diameter the effect was not significant. In attached twigs frozen in field portable freezing chambers, HTEs were recorded at a similar mean temperature (-4.6?±?1.0 °C) to those for detached twigs. Upon lethal intracellular freezing of apple xylem parenchyma cells (XPCs) low-temperature freezing exotherms (LTEs) can be recorded. Low-temperature freezing exotherms determined on detached twigs varied significantly between a winter minimum of -36.9 °C and a summer maximum -12.7 °C. Within the temperature range wherein LTEs were recorded by IDTA in summer (-12.7?±?0.5 to -20.3?±?1.1 °C) various tiny clearly separated discontinuous freezing events could be detected similar to that in other species with contrasting XPC anatomy. These freezing events appeared to be initially located in the primary and only later in the secondary xylem. During the LTE no freezing events in the bark and central pith tissue were recorded. Attached twigs were exposed to various freezing temperatures at which LTEs occur. Even if 60% of XPCs were frost-damaged twigs were able to recuperate and showed full re-growth indicating a high regeneration capacity even after severe frost damage to XPCs.  相似文献   

11.
Tyree MT 《Tree physiology》1988,4(3):195-217
A model is presented for the dynamics of water flow in a single eastern white cedar tree (Thuja occidentalis L.). The model takes into account the spatial and temporal dependence of the evaporative flux from leaves in the crown. It also accounts for the quantitative hydraulic architecture of the tree, i.e., the model characterizes the tree as a branched catena of > 4000 stem segments in which account is taken of the segment length, diameter, hydraulic resistance, and the total area of leaves attached to the segment. Input values needed to run the model are measurements of evaporative flux, hydraulic conductance of stems versus stem diameter, and leaf and stem water storage capacitances. Output parameters are the spatial and temporal characterization of stem and leaf water potentials, stem and leaf water deficits, sap flow rate, and relative sap velocity. The input and output values of the branched catena model are compared and contrasted to that of an unbranched catena model. It is shown that the branched catena model fits independently measured field parameters better than an unbranched catena model. Close correspondence is found between model predictions and field measurements of shoot water potential, pressure gradients in stems, hysteresis in sap velocity between the lower and upper parts of the tree, and diurnal changes in stem and leaf water deficits. This model is discussed in terms of both the hydraulic architecture of trees and the potential application of the model to questions of tree morphology, ecology, physiology and evolution.  相似文献   

12.
I conducted a literature survey to assess the available information on relationships between size--expressed in terms of diameter and dry biomass--and hydraulic efficiency of woody structures at different scales, from stem segments to whole trees. Three data sets were constructed: the first described the relationship between segment diameter and hydraulic conductivity (k(h); kg m s(-1) MPa(-1)) in four species; the second, for the same four species, described the intraspecific trajectories of change in total hydraulic conductance (G; kg s(-1) MPa(-1)) during ontogeny, i.e., from saplings to mature trees, thereby providing a comparison between allometric scaling laws at the scales of segments and whole trees; the third comprised pooled means for nine species that described the interspecific trajectory of change in G with tree size. The scaling coefficients obtained were compared with predictions made with an architectural fractal-like model incorporating tissue-specific hydraulic architecture parameters (West et al. 1999). When data on segment k(h) were examined, the fractal-like model closely predicted the scaling of k(h) with segment diameter in four species. However, the model failed to predict accurately in all species the intraspecific scaling at the branch and whole-tree levels, and consistently overestimated the scaling coefficients. The results suggest that ontogenetic changes in tree size during the life cycle of one tree may result in tradeoffs between optimal hydraulic supply to the existing leaf area and maintenance costs of the supporting xylem tissue. The model of West et al. (1999) may be useful for understanding broad interspecific patterns, but not for understanding more subtle ontogenetic changes.  相似文献   

13.
Tropical moist forests are notable for their richness in tree species. The presence of such a diverse tree flora presents potential problems for scaling up estimates of water use from individual trees to entire stands and for drawing generalizations about physiological regulation of water use in tropical trees. We measured sapwood area or sap flow, or both, in 27 co-occurring canopy species in a Panamanian forest to determine the extent to which relationships between tree size, sapwood area and sap flow were species-specific, or whether they were constrained by universal functional relationships between tree size, conducting xylem area, and water use. For the 24 species in which active xylem area was estimated over a range of size classes, diameter at breast height (DBH) accounted for 98% of the variation in sapwood area and 67% of the variation in sapwood depth when data for all species were combined. The DBH alone also accounted for > or = 90% of the variation in both maximum and total daily sap flux density in the outermost 2 cm of sapwood for all species taken together. Maximum sap flux density measured near the base of the tree occurred at about 1,400 h in the largest trees and 1,130 h in the smallest trees studied, and DBH accounted for 93% of the variation in the time of day at which maximum sap flow occurred. The shared relationship between tree size and time of maximum sap flow at the base of the tree suggests that a common relationship between diurnal stem water storage capacity and tree size existed. These results are consistent with a recent hypothesis that allometric scaling of plant vascular systems, and therefore water use, is universal.  相似文献   

14.
We monitored sap flux density (v) diurnally in nine mature southeastern pine (Pinus spp.) trees with a thermal dissipation probe that spanned the sapwood radius. We found the expected pattern of high v near the cambium and decreasing v with depth toward the center of the tree; however, the pattern was not constant within a day or between trees. Radial profiles of trees were steeper earlier in the day and became less steep later in the day. As a result, time-dependent changes in the shape of the radial profile of v were sometimes correlated with daily changes in evaporative demand. As the radial profile became less steep, the inner xylem contributed relatively more to total tree sap flow than it did earlier in the day. We present a 3-parameter Gaussian function that can be used to describe the radial distribution of v in trees. Parameters in the function represent depth in the xylem from the cambium, maximum v, depth in the xylem where maximum v occurs, and the rate of radial change in v with radial depth (beta). Values of beta varied significantly between trees and with time, and were sometimes correlated with air vapor pressure deficit (D). We hypothesize that this occurred during periods of high transpiration when the water potential gradient became great enough to move water in the inner sapwood despite its probable high hydraulic resistance. We examined discrepancies among estimates of daily water use based on single-point, two-point and multi-point (i.e., every 20 mm in the sapwood) measurements. When radial distribution of v was not considered, a single-point measurement resulted in errors as large as 154% in the estimate of daily water use relative to the estimate obtained from a multi-point measurement. Measuring v at two close sample points (10 and 30 mm) did not improve the estimate; however, estimates derived from v measured at two distant sample points (10 and 70 mm) significantly improved the estimate of daily water use, although errors were as great as 32% in individual trees. The variability in v with depth in the xylem, over time, and between trees indicates that measurements of the radial distribution of v are necessary to accurately estimate water flow in trees with large sapwood areas.  相似文献   

15.
We determined the axial and radial xylem tension gradients in trunks of young Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees. Axial specific conductivity (k(s-a)) and sap flux density (Js) were measured at four consecutive depths within the sapwood at a stem height of 1 m. By definition, at a given position in the bole, Js is a function not only of k(s-a) but also of the driving force for water movement. The Js:k(s-a) ratio was therefore used to estimate axial tension gradients and the radial gradients at a stem height of 1 m were calculated from the differences in axial tension gradients at each depth. Tracheid lumen diameter and tracheid length were used to predict differences in k(s-a) and its divergence from the theoretical k(s-a) determined by the Hagen Poisseuille equation. The ratio of k(s-a) (determined in the laboratory) to Js (measured in the field) varied with depth in the sapwood, resulting in non-uniform axial and radial tension gradients from inner to outer sapwood. Transpiration-induced axial tension gradients were in the range of 0.006-0.01 MPa m(-1) excluding the gravitational tension gradient. At a stem height of 1 m, radial tension gradients were in the range of 0.15-0.25 MPa m(-1) and were lower in the middle sapwood than in the inner or outer sapwood. Axial tension gradients were 44-50% higher in the outer sapwood than in the inner sapwood. At a stem height of 1 m, radial Js, calculated on the basis of radial tension gradients and measured radial specific conductivity (k(s-r)), was about two orders of magnitude smaller than axial Js. Our findings indicate that large radial tension gradients occur in the sapwood and clarify the role played by xylem k(s-a) and k(s-r) in determining in situ partitioning of Js in the axial and radial directions.  相似文献   

16.
We estimated daily use of stored water by Scots pine (Pinus sylvestris L.) trees growing in a temperate climate with the ANAFORE model (ANAlysis of FORest Ecosystems) and compared the simulation results with sap flow measurements. The original model was expanded with a dynamic water flow and storage model that simulates sap flow dynamics in an individual tree. ANAFORE was able to accurately simulate diurnal patterns of measured sap flow under microclimatic conditions that differ from those of the calibration period. Strong relationships were found between stored water use and several tree characteristics (diameter at breast height, sapwood area, leaf area), but not with tree height. Relative to transpiration, stored water use varied over time (between < 1% and 44% of daily transpiration). On days when transpiration was high, trees were more dependent on stored water, indicating that the contribution of internal water to transpiration is not a constant in the water budget of trees.  相似文献   

17.
It is predicted that dryland salinity will affect up to 17 Mha of the Australian landscape by 2050, and therefore, monitoring the health of tree plantings and remnant native vegetation in saline areas is increasingly important. Casuarina glauca Sieber ex Spreng. has considerable salinity tolerance and is commonly planted in areas with a shallow, saline water table. To evaluate the potential of using the nitrogenous composition of xylem sap to assess salinity stress in C. glauca, the responses of trees grown with various soil salinities in a greenhouse were compared with those of trees growing in field plots with different water table depths and groundwater salinities. In the greenhouse, increasing soil salinity led to increased allocation of nitrogen (N) to proline and arginine in both stem and root xylem sap, with coincident decreases in citrulline and asparagine. Although the field plots were ranked as increasingly saline-based on ground water salinity and depth-only the allocation of N to citrulline differed significantly between the field plots. Within each plot, temporal variation in the composition of the xylem sap was related to rainfall, rainfall infiltration and soil salinity. Periods of low rainfall and infiltration and higher soil salinity corresponded with increased allocation of N to proline and arginine in the xylem sap. The allocation of N to citrulline and asparagine increased following rainfall events where rain was calculated to have infiltrated sufficiently to decrease soil salinity. The relationship between nitrogenous composition of the xylem sap of C. glauca and soil salinity indicates that the analysis of xylem sap is an effective method for assessing changes in salinity stress in trees at a particular site over time. However, the composition of the xylem sap proved less useful as a comparative index of salinity stress in trees growing at different sites.  相似文献   

18.
We developed a new microelectrode technique for measuring CO2 concentration ([CO2]) in xylem sap of trees. This technique enabled us to make rapid and continuous measurements of xylem sap [CO2] in situ. In this report, we discuss the methodology and establish the feasibility of the technique. We also describe calibration procedures, temperature sensitivity, field use and other characteristics of the microelectrodes. An example of data collected in the field is provided. Microelectrode calibration was accomplished at constant temperature in air of known [CO2]. When sampling temperature differed from calibration temperature, correction was necessary. We developed an equation to correct for temperatures between 15 and 35 degrees C when calibration was conducted at 25 degrees C. Equations based on Henry's Law were used to convert measured gas phase [CO2] (%) to concentration of all products of CO2 dissolved in sap (mmol l(-1)). We inserted microelectrodes into stems of three tree species to measure diurnal changes in [CO2] in the xylem sap. A diurnal pattern with depression during the day and elevation at night was observed. Mean daily [CO2] ranged from 1.6 to 10.3 mmol l(-1). Microelectrodes were suitable for making diurnal measurements for up to 7 days without recalibration. We also used the microelectrodes to measure [CO2] of soil in situ. Soil [CO2] ranged from 1 to 4% (gas phase), with little diurnal variation.  相似文献   

19.
In tall old forests, limitations to water transport may limit maximum tree height and reduce photosynthesis and carbon sequestration. We evaluated the degree to which tall trees could potentially compensate for hydraulic limitations to water transport by increased use of water stored in xylem. Using sap flux measurements in three tree species of the Pacific Northwest, we showed that reliance on stored water increases with tree size and estimated that use of stored water increases photosynthesis. For Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), water stored in xylem accounted for 20 to 25% of total daily water use in 60-m trees, whereas stored water comprised 7% of daily water use in 15-m trees. For Oregon white oak (Quercus garryana Dougl. ex Hook.), water stored in xylem accounted for 10 to 23% of total daily water use in 25-m trees, whereas stored water comprised 9 to 13% of daily water use in 10-m trees. For ponderosa pine (Pinus ponderosa Dougl. ex Laws.), water stored in xylem accounted for 4 to 20% of total daily water use in 36-m trees, whereas stored water comprised 2 to 4% of daily water use in 12-m trees. In 60-m Douglas-fir trees, we estimated that use of stored water supported 18% more photosynthesis on a daily basis than would occur if no stored water were used, whereas 15-m Douglas-fir trees gained 10% greater daily photosynthesis from use of stored water. We conclude that water storage plays a significant role in the water and carbon economy of tall trees and old forests.  相似文献   

20.
  • ? It has been known for a long time that sectored and integrated patterns of vascular systems exist in different species and even within the same tree, depending on its age and history. However, very few publications consider the topology of the vascular pathways between roots and branches.
  • ? Some results on this important aspect of the vascular system are presented in this paper. They have been obtained with adult maple trees by directly studying the water movement in the stem and root xylem with the heat field deformation (HFD) method for sap flow measurements.
  • ? Multi-point HFD sensors were installed at different heights of a Norway maple tree (Acer platanoides L.) along its stem axis. Single-point HFD sensors were installed in three small lateral roots of another sample maple. Experimental treatments (branch severing) triggered changes in sap movement in the stem and root sapwood.
  • ? The sample trees belong to the group with an integrated transport system (“integrated pipes”), sharing stem space on both sides of the tree to supply two large parts of the crown with water from each root sector. Nevertheless, conducting pathways had their autonomy for axial transport and the pipe model theory describes the vascular system of the studied trees well. Thus, the integration of axial transport in the stem xylem should presumably occur through the cross-grained network of axial vessels.
  •   相似文献   

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