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1.
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   

2.
Trees must respond to many environmental factors during their development, and light is one of the main stimuli regulating tree growth. Thinning of forest stands by selective tree removal is a common tool in forest management that increases light intensity. However, morphological and anatomical adaptations of individual shoots to the new environmental conditions created by thinning are still poorly understood. In this study, we evaluated shoot morphology (shoot length, needle number, projected leaf area) and anatomy (tracheid lumen area, tracheid number, tracheid dimensions, xylem area, potential hydraulic conductivity) in three Norway spruce (Picea abies/L./Karst.) families exposed to different thinning regimes. We compared shoot characteristics of upper-canopy (i.e. sun-exposed) and lower-canopy (i.e. shaded) current-year shoots in a control plot and a plot thinned to 50 % stand density the previous year. One tree per family was chosen in each treatment, and five shoots were taken per canopy position. We found that upper-canopy shoots in both plots had higher values than lower-canopy shoots for all studied parameters, except lumen roundness and tracheid frequency (i.e. tracheid number per xylem area). Thinning had little effect on shoot morphology and anatomy 1 year after thinning, except for small but significant changes in tracheid dimensions. Needles were more sensitive to altered light conditions, as projected leaf area of shoot, needle number and leaf hydraulic conductivity changed after thinning. Differences between upper- and lower-canopy shoots did not seem to be influenced by thinning and were almost the same in both plots. Our results suggest that lower-canopy shoots require several years to modify their morphology and anatomy to new light conditions following thinning. The slow light adaptation of the lower canopy may be of practical importance in forest management: thinned stands may be predisposed to drought stress because newly exposed shoots experience increased illumination and transpiration after thinning.  相似文献   

3.
Ultrasonic emission (UE) testing is used to analyse the vulnerability of xylem to embolism, but the number of UEs often does not sufficiently reflect effects on hydraulic conductivity. We monitored the absolute energy of UE signals in dehydrating xylem samples hypothesizing that (i) conduit diameter is correlated with UE energy and (ii) monitoring of UE energy may enhance the utility of this technique for analysis of xylem vulnerability. Split xylem samples were prepared from trunk wood of Picea abies, and four categories of samples, derived from mature (I: earlywood, II: 30-50% latewood, III: >50% latewood) or juvenile wood (IV: earlywood) were used. Ultrasonic emissions during dehydration were registered and anatomical parameters (tracheid lumen area, number per area) were analysed from cross-sections. Attenuation of UE energy was measured on a dehydrating wood beam by repeated lead breaks. Vulnerability to drought-induced embolism was analysed on dehydrating branches by hydraulic, UE number or UE energy measurements. In split samples, the cumulative number of UEs increased linearly with the number of tracheids per cross-section, and UE energy was positively correlated with the mean lumen area. Ultrasonic emission energies of earlywood samples (I and IV), which showed normally distributed tracheid lumen areas, increased during dehydration, whereas samples with latewood (II and III) exhibited a right-skewed distribution of lumina and UE energies. Ultrasonic emission energy was hardly influenced by moisture content until ～40% moisture loss, and decreased exponentially thereafter. Dehydrating branches showed a 50% loss of conductivity at -3.6 MPa in hydraulic measurements and at -3.9 and -3.5 MPa in UE analysis based on cumulative number or energy of signals, respectively. Ultrasonic emission energy emitted by cavitating conduits is determined by the xylem water potential and by the size of element. Energy patterns during dehydration are thus influenced by the vulnerability to cavitation, conduit size distribution as well as attenuation properties. Measurements of UE energy may be used as an alternative to the number of UEs in vulnerability analysis.  相似文献   

4.
Leaves, the distal section of the soil-plant-atmosphere continuum, exhibit the lowest water potentials in a plant. In contrast to angiosperm leaves, knowledge of the hydraulic architecture of conifer needles is scant. We investigated the hydraulic efficiency and safety of Pinus pinaster needles, comparing different techniques. The xylem hydraulic conductivity (k(s)) and embolism vulnerability (P(50)) of both needle and stem were measured using the cavitron technique. The conductance and vulnerability of whole needles were measured via rehydration kinetics, and Cryo-SEM and 3D X-ray microtomographic observations were used as reference tools to validate physical measurements. The needle xylem of P. pinaster had lower hydraulic efficiency (k(s)?=?2.0?×?10(-4) m(2) MPa(-1) s(-1)) and safety (P(50)?=?-?1.5 MPa) than stem xylem (k(s)?=?7.7?×?10(-4) m(2) MPa(-1) s(-1); P(50)?=?-?3.6 to?-?3.2 MPa). P(50) of whole needles (both extra-vascular and vascular pathways) was?-?0.5 MPa, suggesting that non-vascular tissues were more vulnerable than the xylem. During dehydration to?-?3.5 MPa, collapse and embolism in xylem tracheids, and gap formation in surrounding tissues were observed. However, a discrepancy in hydraulic and acoustic results appeared compared with visualizations, arguing for greater caution with these techniques when applied to needles. Our results indicate that the most distal parts of the water transport pathway are limiting for hydraulics of P. pinaster. Needle tissues exhibit a low hydraulic efficiency and low hydraulic safety, but may also act to buffer short-term water deficits, thus preventing xylem embolism.  相似文献   

5.
Flooding of soil for 55 days altered the rate of growth and stem anatomy of 9‐month‐old Cryptomeria japonica seedlings. Although flooding did not affect height growth it reduced the rate of dry weight increment of seedlings while increasing stem diameter. The reduction in dry weight increment of seedlings resulted largely from decay of roots and, to a lesser extent, from inhibition of growth of roots and needles. The increased diameter growth of flooded seedlings resulted largely from an increase in bark thickness associated with increased phloem production and greater amount of intercellular space. Flooding reduced xylem increment in submerged stems but increased it above the water level because of larger tracheids rather than more tracheids per radial file. Flooding also increased lumen diameters of tracheids, decreased tracheid wall thickness (as a proportion of tracheid diameter), and stimulated formation of axial parenchyma cells in the xylem. Cryptomeria japonica seedlings adapted to flooding by forming adventitious roots, primarily on the original root system and submerged portion of the stem. Such new roots originated in the xylem ray parenchyma. Flooding stimulated ACC synthesis in roots and ethylene production in stems. The role of ethylene in alteration of stem anatomy is discussed.  相似文献   

6.
We examined the effects of increased transpiration demand on xylem hydraulic conductivity and vulnerability to cavitation of mature ponderosa pine (Pinus ponderosa Laws.) by comparing trees growing in contrasting climates. Previous studies determined that trees growing in warm and dry sites (desert) had half the leaf/sapwood area ratio (A(L)/A(S)) and more than twice the transpiration rate of trees growing in cool and moist sites (montane). We predicted that high transpiration rates would be associated with increased specific hydraulic conductivity (K(S)) and increased resistance to xylem cavitation. Desert trees had 19% higher K(S) than montane trees, primarily because of larger tracheid lumen diameters. Predawn water potential and water potential differences between the soil and the shoot were similar for desert and montane trees, suggesting that differences in tracheid anatomy, and therefore K(S), were caused primarily by temperature and evaporative demand, rather than soil drought. Vulnerability to xylem cavitation did not differ between desert and montane populations. A 50% loss in hydraulic conductivity occurred at water potentials between -2.61 and -2.65 MPa, and vulnerability to xylem cavitation did not vary with stem size. Minimum xylem tensions of desert and montane trees did not drop below -2.05 MPa. Foliage turgor loss point did not differ between climate groups and corresponded to mean minimum xylem tensions in the field. In addition to low A(L)/A(S), high K(S) in desert trees may provide a way to increase tree hydraulic conductivity in response to high evaporative demand and prevent xylem tensions from reaching values that cause catastrophic cavitation. In ponderosa pine, the flexible responses of A(L)/A(S) and K(S) to climate may preclude the existence of significant intraspecific variation in the vulnerability of xylem to cavitation.  相似文献   

7.
We studied the effects of elevated temperature and carbon dioxide concentration ([CO(2)]) alone and together on wood anatomy of 20-year-old Scots pine (Pinus sylvestris L.) trees. The study was conducted in 16 closed chambers, providing a factorial combination of two temperature regimes and two CO(2) concentrations (ambient and elevated), with four trees in each treatment. The climate scenario included a doubling of [CO(2)] and a corresponding increase of 2-6 degrees C in temperature at the site depending on the season. Anatomical characteristics analyzed were annual earlywood, latewood and ring widths, intra-ring wood densities (earlywood, latewood and mean wood density), tracheid width, length, wall thickness, lumen diameter, wall thickness:lumen diameter ratio and mass per unit length (coarseness), and numbers of rays, resin canals and tracheids per xylem cross-sectional area. Elevated [CO(2)] increased ring width in four of six treatment years; earlywood width increased in the first two years and latewood width in the third year. Tracheid walls in both the earlywood and latewood tended to become thicker over the 6-year treatment period when temperature or [CO(2)] was elevated alone, whereas in the combined treatment they tended to become thinner relative to the tracheids of trees grown under ambient conditions. Latewood tracheid lumen diameters were larger in all the treatments relative to ambient conditions over the 6-year period, whereas lumen diameters in earlywood increased only in response to elevated [CO(2)] and were 3-6% smaller in the treatments with elevated temperature than in ambient conditions. Tracheid width, length and coarseness were greater in trees grown in elevated than in ambient temperature. The number of resin canals per mm(2) decreased in the elevated [CO(2)] treatment and increased in the elevated temperature treatments relative to ambient conditions. The treatments decreased the number of rays and tracheids per mm(2) of cross-sectional area, the greatest decrease occurring in the elevated [CO(2)] treatment. It seemed that xylem anatomy was affected more by elevated temperature than by elevated [CO(2)] and that the effects of temperature were confined to the earlywood.  相似文献   

8.
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   

9.
Roots from healthy and diseased mature ponderosa pine, Pinus ponderosa Laws., trees were excavated from a site near Burns, Oregon. The diseased trees were infected with black-stain root disease, Leptographium wageneri Kendrick, or annosus root disease, Heterobasidion annosum (Fr.) Bref., or both. Axial hydraulic conductivity of the roots was measured under a positive head pressure of 5 kPa, and the conducting area was stained with safranin dye to determine specific conductivity (k(s)). In diseased roots, only 8-12% of the cross-sectional xylem area conducted water. Resin-soaked xylem completely restricted water transport and accounted for 13-16% of the loss in conducting area. In roots with black-stain root disease, 17% of the loss in conducting area was associated with unstained xylem, possibly resulting from occlusions or embolisms. Based on the entire cross-sectional area of infected roots, the k(s) of roots infected with black-stain root disease was 4.6% of that for healthy roots, whereas the k(s) of roots infected with annosus root disease was 2.6% of that for healthy roots. Although these low values were partly the result of the presence of a large number of diseased roots (72%) with no conducting xylem, the k(s) of functional xylem of diseased roots was only 33% of that for healthy roots. The low k(s) values of functional xylem in diseased roots may be caused by fungus induced occlusions preceding cavitation and embolism of tracheids. The k(s) of disease-free roots from diseased trees was only 70% of that for healthy roots from healthy trees. The disease-free roots had the same mean tracheid diameter and tissue density as the healthy roots, suggesting that the lower k(s) in disease-free roots of diseased trees may also have been caused by partial xylary occlusions.  相似文献   

10.
The effect of water-stress conditioning on water relations and histological features ofPinus thunbergii Parl. inoculated with avirulent isolate ofBursaphelenchus xylophilus (Steiner and Buhrer) Nickle, pine wood nematode, were investigated. Pines were kept under 8 days cycle of severe water stress. One-half of the water-stressed pines died as a result of infection by avirulent pine wood nematode and water stress tended to induce increased susceptibility and/or decreased resistance of pines to avirulent pine wood nematode. In dead pines, the water conducting function of xylem was lost, and all of the parenchyma cells died. In surviving pines, the xylem hydraulic conductivity and the xylem water content were significantly reduced (12 to 23% and 77 to 83%, respectively) compared to controls. Safranin dye perfusion of excised axis stem segments indicated that the water conductance was limited to the very narrow peripheral area of xylem. Embolism caused by cavitation in the tracheids occurred in the central part of xylem and in that dysfunctional region of the xylem the axial parenchyma cells surrounding the epithelial cells, and ray parenchyma cells partly degenerated but the epithelial cells survived. The disruption of tracheid shape observed in surviving pines indicates that avirulent pine wood nematode temporarily disturbed cell division of the cambium. Considering the differences in responses between dead pines and surviving pines after inoculation with avirulent pine wood nematode, the death of water-stressed pines apparently resulted from death of cells, in particular the vascular cambium and the loss of xylem hydraulic function by cavitation.  相似文献   

11.
Aumann CA  Ford ED 《Tree physiology》2006,26(3):285-301
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   

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.
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   

14.
Phenotypic plasticity in needle morphology with increasing tree size and age was investigated by comparing four age classes of red spruce (Picea rubens Sarg.) ranging from juvenile (3-12 years old) to mature (over 100 years old). With increase in tree age there were significant increases in leaf mass per unit area (LMA), mesophyll and vascular bundle area as a percentage of total needle cross-sectional area, and stomatal density. Within the vascular bundle, both xylem cross-sectional area and tracheid lumen area increased significantly, whereas air space as a percentage of total cross-sectional area decreased. These morphological changes were associated with a significant decrease in photosynthetic capacity and stomatal conductance, and an increase in (13)C enrichment. Although both photosynthetic capacity and whole-tree conductance decreased significantly between age classes 3 and 12 years, they did not differ between age classes 53 and 127 years, even though needle (13)C enrichment was significantly greater in the 127-year age class. Thus there appear to be compensatory mechanisms that maintain photosynthetic capacity as trees increase in size and vascular complexity, which in red spruce and other species, may affect leaf hydraulic conductance. Although increased LMA may contribute to reduced photosynthetic capacity in red spruce, similar relationships are not seen in other conifers.  相似文献   

15.
We investigated the impact of drought on the physiology of 41-year-old Scots pine (Pinus sylvestris L.) in central Scotland. Measurements were made of the seasonal course of transpiration, canopy stomatal conductance, needle water potential, xylem water content, soil-to-needle hydraulic resistance, and growth. Comparison was made between drought-treated plots and those receiving average precipitation. In response to drought, transpiration rate declined once volumetric water content (VWC) over the top 20 cm of soil reached a threshold value of 12%. Thereafter, transpiration was a near linear function of soil water content. As the soil water deficit developed, the hydraulic resistance between soil and needles increased by a factor of three as predawn needle water potential declined from -0.54 to -0.71 MPa. A small but significant increase in xylem embolism was detected in 1-year-old shoots. Stomatal control of transpiration prevented needle water potential from declining below -1.5 MPa. Basal area, and shoot and needle growth were significantly reduced in the drought treatment. In the year following the drought, canopy stomatal conductance and soil-to-needle hydraulic resistance recovered. Current-year needle extension recovered, but a significant reduction in basal area increment was evident one year after the drought. The results suggest that, in response to soil water deficit, mature Scots pine closes its stomata sufficiently to prevent the development of substantial xylem embolism. Reduced growth in the year after a severe soil water deficit is most likely to be the result of reduced assimilation in the year of the drought, rather than to any residual embolism carried over from one year to the next.  相似文献   

16.
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   

17.
The model of West, Brown and Enquist (1999) shows that hydraulic resistance in trees can be independent of path length, provided that vascular conduits widen sufficiently from tree top to base. We demonstrate that this result does not depend theoretically on branching architecture or cross-sectional conductive area of the stem. Previous studies have shown that pit membrane resistance, encountered when water moves between either tracheids or vessels, accounts for up to 60% of the total resistance in stem segments. When pit membrane resistance, which is neglected by most whole-tree hydraulic models, was incorporated in hydraulic models in three different ways, the near invariance of hydraulic resistance was preserved. If relative pit resistance was independent of tracheid size or if tracheid dimensions were scaled to minimize wood resistivity, the minimum conduit taper required for path length independence equaled that in the original model of West et al. (1999). Under the most realistic model, in which relative pit resistance increased with tracheid radius, this value was doubled. Such taper is not possible within the typical size range of tracheids over the entire length of moderately tall trees, but it might be possible for vessel-bearing trees. Preliminary results indicated that although tracheid radius in the outer growth ring initially increased basipetally from the top of an 18-m tall Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), it stabilized at mid-trunk. Also, conduit taper was not constant in this species, violating a key assumption of the model of West et al. (1999), on which the invariance of hydraulic resistance depends.  相似文献   

18.
Ladjal M  Huc R  Ducrey M 《Tree physiology》2005,25(9):1109-1117
We studied hydraulic traits of young plants of the Mediterranean cedar species Cedrus atlantica (Endl.) G. Manetti ex Carrière (Luberon, France), C. brevifolia (Hook. f.) Henry (Cyprus), C. libani A. Rich (Hadeth El Jebbe, Lebanon) and C. libani (Armut Alani, Turkey). With an optimum water supply, no major differences were observed among species or provenances in either stem hydraulic conductivity (Ks) or leaf specific conductivity (Kl) measured on the main shoot. A moderate soil drought applied for 10 weeks induced marked acclimation through a reduction in Ks, particularly in the Lebanese provenance of C. libani, and a decrease in tracheid lumen size in all species. Cedrus atlantica, which had the smallest tracheids, was the species most vulnerable to embolism: a 50% loss in hydraulic conductivity (PsiPLC50) occurred at a water potential of -4.4 MPa in the well-watered treatment, and at -6.0 MPa in the moderate drought treatment. In the other species, PsiPLC50 was unaffected by moderate soil drought, and only declined sharply at water potentials between -6.4 and -7.5 MPa in both irrigation treatments. During severe drought, Ks of twigs and stomatal conductance (g(s)) were measured simultaneously as leaf water potential declined. For all species, lower vulnerability to embolism based on loss of Ks was recorded on current-year twigs. The threshold for stomatal closure (10% of maximum g(s)) was reached at a predawn water potential (Psi(pd)) of -2.5 MPa in C. atlantica (Luberon) and at -3.1 MPa in C. libani (Lebanon), whereas the other provenance and species had intermediate Psi(pd) values. Cedrus brevifolia, with a Psi(pd) (-3.0 MPa) close to that of C. libani (Lebanon), had the highest stomatal conductance of the study species. The importance of a margin of safety between water potential causing stomatal closure and that causing xylem embolism induction is discussed.  相似文献   

19.
Domec JC  Pruyn ML 《Tree physiology》2008,28(10):1493-1504
Effects of trunk girdling on seasonal patterns of xylem water status, water transport and woody tissue metabolic properties were investigated in ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) trees. At the onset of summer, there was a sharp decrease in stomatal conductance (g(s)) in girdled trees followed by a full recovery after the first major rainfall in September. Eliminating the root as a carbohydrate sink by girdling induced a rapid reversible reduction in g(s). Respiratory potential (a laboratory measure of tissue-level respiration) increased above the girdle (branches and upper trunk) and decreased below the girdle (lower trunk and roots) relative to control trees during the growing season, but the effect was reversed after the first major rainfall. The increase in branch respiratory potential induced by girdling suggests that the decrease in g(s) was caused by the accumulation of carbohydrates above the girdle, which is consistent with an observed increase in leaf mass per area in the girdled trees. Trunk girdling did not affect native xylem embolism or xylem conductivity. Both treated and control trunks experienced loss of xylem conductivity ranging from 10% in spring to 30% in summer. Girdling reduced xylem growth and sapwood to leaf area ratio, which in turn reduced branch leaf specific conductivity (LSC). The girdling-induced reductions in g(s) and transpiration were associated with a decrease in leaf hydraulic conductance. Two years after girdling, when root-to-shoot phloem continuity had been restored, girdled trees had a reduced density of new wood, which increased xylem conductivity and whole-tree LSC, but also vulnerability to embolism.  相似文献   

20.
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   

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