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1.
The model of West, Brown and Enquist showed that total hydraulic resistance in trees can be independent of path length, provided that vascular conduits taper sufficiently. This model assumes that the tree branch network is volume-filling, so that segment lengths increase exponentially from tree top to base. We show that partial buffering of hydraulic resistance from path-length effects can occur even for moderate tapering, and that this effect is stronger when segment lengths are fixed. Still needed are measurements of tracheary size and hydraulic resistance designed to test this model, which shows how hydraulic limitation of tree height growth may be mitigated. 相似文献
2.
A tracheid-level model was used to quantify the effects of differences in wood microstructure between coastal and interior Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. menziesii and var. glauca) wood on larger scale properties like hydraulic conductivity. The model showed that tracheid length, the ease of flow through a bordered pit and effective tracheid diameter can all limit maximum hydraulic conductivity. Among the model parameters tested, increasing bordered pit conductivity and tracheid length resulted in the greatest increase in maximum conductivity in both the inland and coastal ecotypes. A sensitivity analysis of the uncertainty between parameters governing flow through the bordered pit and air-seeding potential showed that, although decreased pit flow resistance increased maximum hydraulic conductivity, increased cavitation led to lower conductivity over time. The benefits of increasing the number of bordered pits depended on the intensity of the meteorological driving function: in drier environmental conditions, wood with fewer pits was more conductive over time than wood with more pits. Switching the bordered pit characteristics between coastal and interior wood indicated that the conductivity time course of coastal and interior wood was primarily governed by differences in the number of bordered pits and not differences in tracheid dimensions. The rate at which tracheids refilled had little effect on the conductivity time course of either coastal or interior wood during the first two summers when the wood was highly saturated, but had a marked influence in subsequent years once the cavitation profile stabilized. Our work highlights the need for more empirical work on bordered pits to determine whether variation in their number and properties is related to changing environmental conditions. In addition, a detailed simulation model of a bordered pit is needed to understand how variation in pit properties affects the relationship between ease of flow through a bordered pit and its potential for facilitating air-seeding. 相似文献
3.
人工兴安落叶松次生木质部的解剖学研究 总被引：1，自引：0，他引：1
运用木材解剖图像分析系统和显微照相的方法对人工兴安落叶松次生木质部的解剖结构进行研究,结果表明:落叶松具正常树脂道和受伤树脂道两种类型,前者常见于晚材。落叶松生长轮内的早晚材在干和枝内急变,在根内缓变。早材管胞呈六边形至多边形,胞壁常见单列具缘纹孔,偶见对列具缘纹孔;晚材管胞多呈矩形,胞壁鲜见具缘纹孔,通常为单列具缘纹孔。落叶松木射线同时具有单列木射线和纺锤形木射线两种类型,纺锤形木射线中仅含一枚纵行树脂道。纵行管胞与木射线交叉形成的纹孔场为云杉型。从根到干再到枝,管胞逐渐细化,管胞长度逐渐减小,木射线分布由密到疏。 相似文献
4.
Roman Gebauer Roman Plichta Emilie Bednářová Jiří Foit Václav Čermák Josef Urban 《European Journal of Forest Research》2018,137(1):57-67
While needles represent a proportionally large fraction of whole-plant hydraulic resistance, no studies to date have investigated how source–sink disturbances affect needle xylem structure. In this study, we evaluated structural changes in xylem in current-year needles of Scots pine 227 and 411 days after stem girdling (hereafter referred to as DAG). Maximum and minimum tracheid lumen diameters and therefore also the size of tracheid lumen areas increased in needles 227 DAG compared to control needles. In contrast, tracheid dimensions were similar in needles 411 DAG as in the control needles, but smaller xylem area and lower number of tracheids resulted in the lower theoretical needle hydraulic conductivity of those needles. Several needle xylem parameters were intercorrelated in both control and girdled trees. These observed changes provide a new understanding of the processes that occur following a source–sink disturbance. Considering anatomical parameters such as the number of tracheids, tracheid dimension, or needle xylem area, which are rarely described in physiological studies, could be helpful, for example, in understanding to tree hydraulic systems or for modeling gas exchange. Finally, empirical equations were developed to calculate needle theoretical hydraulic conductivity and the number of tracheids in needles using an easily measurable parameter of needle xylem area. 相似文献
5.
Hydraulic and mechanical properties of young Norway spruce clones related to growth and wood structure 总被引：3，自引：0，他引：3
Stem segments of eight five-year-old Norway spruce (Picea abies (L.) Karst.) clones differing in growth characteristics were tested for maximum specific hydraulic conductivity (k(s100)), vulnerability to cavitation and behavior under mechanical stress. The vulnerability of the clones to cavitation was assessed by measuring the applied air pressure required to cause 12 and 50% loss of conductivity (Psi(12), Psi(50)) and the percent loss of conductivity at 4 MPa applied air pressure (PLC(4MPa)). The bending strength and stiffness and the axial compression strength and stiffness of the same stem segments were measured to characterize wood mechanical properties. Growth ring width, wood density, latewood percentage, lumen diameter, cell wall thickness, tracheid length and pit dimensions of earlywood cells, spiral grain and microfibril angles were examined to identify structure-function relationships. High k(s100) was strongly and positively related to spiral grain angle, which corresponded positively to tracheid length and pit dimensions. Spiral grain may reduce flow resistance of the bordered pits of the first earlywood tracheids, which are characterized by rounded tips and an equal distribution of pits along the entire length. Wood density was unrelated to hydraulic vulnerability parameters. Traits associated with higher hydraulic vulnerability were long tracheids, high latewood percentage and thick earlywood cell walls. The positive relationship between earlywood cell wall thickness and vulnerability to cavitation suggest that air seeding through the margo of bordered pits may occur in earlywood. There was a positive phenotypic and genotypic relationship between k(s100) and PLC(4MPa), and both parameters were positively related to tree growth rate. Variability in mechanical properties depended mostly on wood density, but also on the amount of compression wood. Accordingly, hydraulic conductivity and mechanical strength or stiffness showed no tradeoff. 相似文献
6.
We evaluated whether patterns in hydraulic architecture increase transport efficiency. Five patterns are identified: area-preserving branching; variable trunk versus twig sap velocity; distally decreasing leaf specific conductivity (K(L)) and conduit diameter; and a decline in leaf specific conductance (k(L)) of the entire plant with maturation. These patterns coexist in innumerable combinations depending on the ratio of distal/proximal conduit number (F). The model of West and colleagues does not account for this diversity, in part by specifying F = 1 and requiring a specific conduit taper derived from the incorrect premise that k(L) is constant with plant size. We used Murray's law to identify the conduit taper that maximizes k(L)for a given vascular investment. Optimal taper requires the ratio of distal/proximal conduit diameter to equal the ratio of distal/proximal K(L). The smaller these ratios, the greater the k(L). Smaller ratios are achieved by an increase in F. Conductivity and diameter ratios < 1 and F >/= 1 in plants are therefore consistent with maximizing conducting efficiency. However, the benefit of increasing F requires area-increasing conduit branching, potentially leading to mechanical instability of trees. This trade-off may explain why tree stems were relatively inefficient with F near 1 and limited conduit taper compared with vine stems or compound leaves with F > 1 and greater taper. Within trees, the anatomies of a coniferous and a diffuse-porous species were less efficient than that of a ring-porous species, presumably because the latter allows conduit area to increase distally without also increasing total xylem area. This is consistent with decelerating sap velocities from trunk to twigs in ring-porous trees versus accelerating velocities in other types. In general, the observed architectural patterns are consistent with the maximization of transport efficiency operating within mechanical constraints. 相似文献
7.
We tested the effect of soil compaction on Norway spruce seedlings in terms of the size and theoretical volume flow rate of
the tracheids. The results show that soil pressure limits growth in the diameter of the lumens of tracheids in all parts of
seedlings studied. The tracheids of the roots with primary xylem had larger lumens than those of the roots and shoots with
secondary xylem in both unloaded and loaded seedlings. This corresponds to the higher cumulative theoretical volume flow rate
of the tracheids from roots with primary xylem than those from roots and shoots with secondary xylem. Although the volume
flow rate of tracheids, according to the Hagen-Poiseuille law, was directly proportional to the quadratic power of the capillary
diameter (tracheid lumen), the cumulative curve of the theoretical hydraulic volume flow rate was higher or relatively comparable
in loaded seedlings. An explanation for these findings is that there were higher gradients of water potential values in roots
and leaves in loaded seedlings because the lengths of the conductive pathways were 27% shorter than in unloaded seedlings.
We hypothesise that trees have adapted to different stresses by shortening their conductive pathways to maintain a transpiration
rate similar to that of non-stressed trees. These results concerning the impact of soil compaction on tracheid diameter and
volume flow rate improve our understanding of the growth and functioning of different conifer organs and the mechanisms underlying
the efficiency of water transport through the root xylem to the shoot. 相似文献
8.
9.
Summary The cross-sectional view of pitting between various cell types inPinus banksiana Lamb. was studied at the ultrastructural level. Cell types inPinus banksiana include longitudinal tracheids, ray tracheids, ray parenchyma cells, buffer cells and epithelial cells. Two common characteristic features of bordered pit-pairs between longitudinal tracheids are an initial pit border and a thickened torus at the center of the pit membrane. The shape and size of the pit border and torus of bordered pit-pairs between two compression wood cells, and between the last-formed latewood longitudinal tracheid and first-formed earlywood longitudinal tracheid were different from those in the earlywood and latewood longitudinal tracheids. The pit border on the ray tracheid side varied in size and shape due to wall dentation. No initial pit border was found on the pit border of the ray tracheid side. The shape of bordered pit-pairs between two ray tracheids varied considerably due to irregularity of the dentate cell wall. The size of bordered pit-pairs in longitudinal tracheids was between 16 m to 20 m, which was twice the diameter of bordered pit-pairs in ray tracheids. Bordered pitpairs at the end wall of two ray tracheids appeared to be the smallest at 5 m, Pit aspiration occurred in the bordered pit-pairs with or without a torus. In the heartwood zone, some half-borders pit-pairs between tracheary and ray parenchyma cells showed an additional secondary wall on the ray parenchyma cell side. Plasmodesmata were found in the half-bordered pit-pairs as well in the simple pit-pairs. Blind pits were observed between a ray tracheid and a longitudinal tracheid. Bordered pit-pairs between two buffer cells were also observed. The possible functions of buffer cells were discussed.Use of transmission electron microscope provided by the Science Instrumentation Lab, Lakehead University and the technical assistance provided by Mr. A. MacKenzie, Director of Science Instrumentation Lab are gratefully appreciated 相似文献
10.
We tested the hypotheses that freezing-induced embolism is related to conduit diameter, and that conifers and angiosperms with conduits of equivalent diameter will exhibit similar losses of hydraulic conductivity in response to freezing. We surveyed the freeze-thaw response of conifers with a broad range of tracheid diameters by subjecting wood segments (root, stem and trunk wood) to a freeze-thaw cycle at -0.5 MPa in a centrifuge. Embolism increased as mean tracheid diameter exceeded 30 microm. Tracheids with a critical diameter greater than 43 microm were calculated to embolize in response to freezing and thawing at a xylem pressure of -0.5 MPa. To confirm that freezing-induced embolism is a function of conduit air content, we air-saturated stems of Abies lasiocarpa (Hook.) Nutt. (mean conduit diameter 13.7 +/- 0.7 microm) by pressurizing them 1 to 60 times above atmospheric pressure, prior to freezing and thawing. The air saturation method simulated the effect of increased tracheid size because the degree of super-saturation is proportional to a tracheid volume holding an equivalent amount of dissolved air at ambient pressure. Embolism increased when the dissolved air content was equivalent to a mean tracheid diameter of 30 microm at ambient air pressure. Our centrifuge and air-saturation data show that conifers are as vulnerable to freeze-thaw embolism as angiosperms with equal conduit diameter. We suggest that the hydraulic conductivity of conifer wood is maximized by increasing tracheid diameters in locations where freezing is rare. Conversely, the narrowing of tracheid diameters protects against freezing-induced embolism in cold climates. 相似文献
11.
油松管胞形态特征的变异 总被引：13，自引：0，他引：13
本文研究分析了山西中条山产地油松管胞形态特征的变异。管胞长度自髓心向外,首先迅速增加,13年后管胞长度增加缓慢,20年后保持相对稳定。管胞长度沿树干主轴自基部向上逐渐增加,5.3m高处最长,然后向上变短,树冠区域管胞长度最短。形成层原始细胞长度随着原始细胞年增大,开始递增,达到最大值后又递减。管胞直径、胞壁厚度自髓心向外增加。管胞直径轴向变化由树干基部开始向上增大,然后又减小。管胞长宽比、壁腔比的径向变异与管胞长度的径向变异模式相似。生长轮内管胞长度从早材到晚材,开始减小,然后增加,最小值位于早晚材过渡处。 相似文献
12.
Compression wood (CW), which is formed on the underside of conifer branches, exhibits a lower specific hydraulic conductivity (k(s)) compared with normal wood. However, the first-formed tracheids of an annual ring on the underside of a conifer branch often share several properties with normal tracheids, e.g., thin cell walls and angular cross sections. These first-formed tracheids appear bright when observed by the naked eye and are therefore called light bands (LB). In this study, hydraulic and related anatomical properties of LBs were characterized and compared with typical CW and opposite wood (OW). Measurements were made on branches of Norway spruce (Picea abies (L.) Karst.). Specific hydraulic conductivity was measured with fine cannulas connected to microlitre syringes. Micro- and ultrastructural analysis were performed on transverse and radial longitudinal sections by light and scanning electron microscopy. Xylem areas containing both typical CW and LBs had a k(s) 51.5% that of OW (7.95 +/- 0.97 m(2) s(-1) MPa(-1) x 10(-4)), whereas k(s) of pure CW was only 26.7% that of OW. The k(s) of LBs (6.38 +/- 0.97 m(2) s(-1) MPa(-1) x 10(-4); 80.3% of OW) was estimated from these k(s) values because the cannulas were too wide to measure the k(s) of LBs directly. Mean lumen area of first-formed tracheids on the underside of branches was 65.7% that of first-formed tracheids in OW and about three times that of CW. Light-band tracheids exhibited a bordered pit frequency of 42.7 +/- 1.3 pits mm(-1), which was three times that in CW and 1.6 times that in OW. Bordered pit apertures in LB tracheids (9.15 +/- 0.60 microm(2)) were 1.7 times wider than those in CW and similar in aperture to those in OW. The high k(s) of LBs was correlated with their wide tracheid lumina, high pit frequency and wide pit apertures. We therefore suggest that LBs have a primarily hydraulic function within the mechanically optimized CW region. This might be important for supplying water to living tissues on the underside of branches, as well as to other distal areas along water transport pathways following the spiral grain of wood. 相似文献
13.
Summary The effects of air-drying and solvent-drying on the sapwood of Abies grandis have been investigated by a new method for the determination of the size and number of conducting tracheid lumina and pit membrane pores which involves the measurement of gaseous permeability at various mean pressures. Both earlywood and latewood tracheids (83% of the total) were found to be conducting in solvent-dried wood, but in air-dried wood only latewood tracheids (32% of the total) were conducting. In solvent-dried wood there were on average 27,000 pit membrane pores per conducting tracheid compared with only 600 in air-dried wood. In both, the average pit membrane pore radius was about 0.1 m.Liquid permeabilities have been predicted from the calculated radii and numbers. The liquid permeability of solvent-dried wood was 31 times greater than that of air-dried wood in which the lumina were responsible for 13% of the total resistance to flow. The lumina were responsible for 39% of the resistance in solvent-dried wood and it is suggested that in first-formed earlywood the lumina may cause more than half the total resistance.A new method is described for the cleaning of direct carbon replicas of wood. In this the cellulose is removed by cellulase instead of sulphuric acid, and no wax backing is required. This provides much cleaner replicas. Electron micrographs have been obtained of both earlywood and latewood dried by the two methods.The authors wish to thank Mr. A. R. Sayers for preparing the computer programme used in this work, Dr. R. Ph. C. Johnson for his help and advice regarding the electron microscopy and Professors Matthews and Weatherley for their advice and encouragement. 相似文献
14.
Dynamics of water storage in mature subalpine Picea abies: temporal and spatial patterns of change in stem radius 总被引：1，自引：0，他引：1
Internal water reserves in bark and foliage of trees contribute to transpiration (T) and play an essential role in optimizing water transport by buffering extreme peaks of water consumption. We examined patterns of stem shrinkage and their relationship to tree water dynamics. We measured fluctuations in root radius and stem radius at different stem heights, T of twigs at the top of the crown and sap flow velocities in stem sections of mature subalpine Norway spruce (Picea abies (L.) Karst.) trees over 2 years. The output of each sensor was coupled by physical functions to a mechanistic flow and storage model of tree water relations. The data verified the model-predicted lag in water storage depletion in response to the onset of transpiration and the lag increased with increasing distance from the crown periphery. Between the crown and stem base, the delay ranged from a few minutes to several hours, depending on microclimatic conditions and tree water status. Stem volume changes were proportional to the amount of water exchanged between the elastic tissues of the bark and the rigid xylem, indicating that the "peristaltic" wave of stem contraction along the flow path represented depletion of water stored in bark. On a daily basis, stems lost between 0.2 and 0.5% of their volume as a result of bark dehydration, corresponding to about 2 to 5 l of water. This water contributed directly to T. According to the model based on hydraulic principles, there are three main components underlying the dynamics of water storage depletion: flow resistance, storage capacities of needles and bark, and T of each tree section. The resistances and capacities were proportional to the response delay, whereas T in the lower parts of the tree was inversely proportional. The pattern of T within the crown depended on water intercepted by the branches. Because of these weather-dependent factors, there was no time constant for the response delay along the flow path. Nevertheless, the upper crown and the root section tended to have longer response delays per meter of flow path than the stem. The diurnal course of stem radius fluctuations represents the sum of all external and internal conditions affecting tree water relations; stem radius fluctuations, therefore, provide a sensitive measure of tree water status. 相似文献
15.
J. A. Petty 《Wood Science and Technology》1987,21(1):43-48
Summary It is shown that to calculate the rate of gaseous diffusion through an isolated capillary of radius a an end correction of a/2 must be added to the physical length of the capillary. This correction is applicable to diffusion through pit apertures in conifer wood. Mutual interference between diffusion through closely spaced capillaries reduces the end correction. Two approaches to the calculation of this effect are discussed. The pores in many bordered pit membranes are so closely spaced that the end correction becomes negligible. It is shown theoretically that when the end correction is taken into account the tracheid lumen/cell wall pathway provides the path of least resistance for transverse moisture diffusion through conifer wood. 相似文献
16.
针对树材气体渗透性与气体在管胞中的流动特性和规律紧密相关，研究管胞的气体渗透流阻，可以获得关于针叶树材气体渗透性更深入的认识。本文从针叶树材管胞解剖结构出发，运用流体力学理论，导出了管胞3个纹理方向渗透气体流阻的数学表达式，并由此计算了针叶树材管胞的气体渗透系数。计算结果表明，管胞流阻可以描述气体渗透阻力的分布情况，反映气体渗透性在3个纹理方向的差异，在木材细胞层次揭示针叶树材气体渗透性的机制。 相似文献
17.
The progress of xylem formation in Norway spruce (Picea abies (L.) Karst.) was measured during one growing season in southern Finland. Stem radius was monitored continuously with band dendrometers, and the formation of new tracheids was determined by examination of small increment cores taken twice weekly. Tracheid production started in June and ceased in August. Xylem formation was fastest in early July, when 0.75-1.25 new tracheids were formed per day. The rate of xylem formation was significantly correlated with mean daily temperature. Synchronous fluctuations in tracheid and lumen diameters were observed at the same relative positions within each annual ring, but no relationship existed between the diameters and weather variables. The timing of changes in stem radius differed from the timing of actual xylem formation. Stem radius increased in April and May, and the fastest daily increments were recorded in June. Increases in stem radius slowed in July, but small increases were measured more than a month after xylem formation had ceased. Daily changes in stem radius were correlated with daily precipitation, reflecting changes in stem water content. Therefore, dendrometers are of dubious value for measuring the timing of actual xylem formation. Small increment cores proved to be useful in assessing actual xylem formation, but the method is laborious. 相似文献
18.
Summary Individual tracheid lengths were measured in macerated tissue samples of Pinus merkussii Jungh. and de Vries, P. kesiya Royle ex Gordon (syn. P. khasya Royle) and P. patula Schiede and Deppe. The sample trees were selected from stands of different ages, treatments and sites in central Africa.A sampling unit of 50 tracheids, measured for convenience on two microscope slides, was chosen on the basis of an earlier study. The variation in individual tracheid length and the variation of within-slide variance was studied by means of analysis of variance and variance components. This indicated the nunber of samples required in the different sampling categories to give maximum sampling efficiency. Sampling categories included both random factors (individual tracheids, number of slides, radii in discs) and fixed factors (annual rings, tree classes, thinning treatments, etc.).While large effects were attributable to fixed factors more than half the total variation was explained by random factors. The effect of the number of slides was rarely important and two slides need be used only to facilitate the location of the required number of tracheids and to provide a check on operator consistency. The sample unit of 50 tracheids can detect differences in tracheid length of 0.20 ... 0.34 mm thus confirming the routine used in this laboratory as a practical and meaningful procedure.Laboratory work was carried out in Kitwe, computing at Oxford. Paper published with permission of Professor of Forest Science, Oxford and Director, Agricultural Research Council of Zambia. 相似文献
19.
Yoshio Kijidani Takahiro Nagai Tomohiko Suwashita Taku Tsuyama 《Journal of Wood Science》2017,63(4):315-321
The role of phytohormones in tracheid formation remains unclear in conifers. In this study, to obtain information on the role of auxin (IAA) and gibberellin A4 (GA4), we examined seasonal variation in the amount of phytohormones in cambial-region tissues and tracheid formation of sugi cultivar planted in a Nelder plot with different tree densities. We demonstrated that the amount of IAA was positively correlated with the number of tracheids formed in early and mid-season, but not in late season, and had no relation to tracheid differentiation. Crown length and height at the crown base had a positive and negative effect, respectively, on IAA amounts in early and mid-season, but not in late season. Height-to-diameter ratio was negatively correlated with IAA amounts in early and mid-season, but not in late season. Sugi trees with wider spacing continued tracheid formation in late season with smaller amounts of IAA, although the trees with narrower spacing ceased tracheid formation with larger amounts of IAA. Cambial growth cessation in late season might be controlled not by IAA amount, but by short-day-induced insensitivity to IAA. GA4 had no relation with the indexes of growth traits or tracheid formation. 相似文献
20.
Mencuccini M 《Tree physiology》2002,22(8):553-565
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