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1.
NIKLAS  KARL J. 《Annals of botany》1997,79(3):265-272
Variations in the density and stiffness (Young's elastic modulus)of fresh wood samples drawn from different parts of the threemain trunks of a 32-year-old black locust tree,Robinia pseudoacacia(measuring 19.8 m at its highest point), were studied to determinewhether tree ontogeny can achieve a constant safety factor againstmechanical failure. Based on the properties of isolated woodsamples, the fresh density of sapwood decreased along radialtransects from bark to pith, while that of progressively olderheartwood samples increased, on average, towards the centreof each of the three trunks. Along the same radial transects,the Young's elastic modulus of sap- and heartwood increased.In terms of longitudinal changes in wood properties, mean woodmoduli (averages of sap- and heartwood samples) increased, onaverage, towards the base of each of the three trunks of thetree. However, the mean fresh densities of wood samples increasedtowards the top and the bottom of each trunk and were lowestroughly near trunk mid-length. The mean density-specific stiffness(the quotient of Young's modulus and fresh density) of woodwas thus lower toward the top and the bottom of the trunks andhighest near trunk mid-length. Mean values of fresh wood density-specificstiffness were used to estimate the critical buckling heightsfor sections of the trunks differing in diameter and age. Theseestimates indicated that ontogenetic variation in the physicalproperties and relative amounts of sap- and heartwood in trunkscould maintain a constant factor of safety (approximately equalto 2) as a sapling grows in height and girth into a mature tree.This expectation was supported by data from 16 black locusttrees differing in height and diameter at breast height (DBH). Wood; elastic properties; tree height; biomechanics  相似文献   

2.
The formation and vertical distribution of sapwood and heartwood were studied with a 45-year-old Cryptomeria japonica D. Don. The tree was grown at a plantation with 1.5 m × 3.0 m spacing near Miao-Li, Taiwan and was felled on 27 February 1992. The thickness of sapwood and heartwood was expressed by a ring count and a linear measurement. The east-west (E-W) wood strips were collected from 0.3 m above ground upwards to the top of the tree at 2.5 m intervals. The sapwood thicknesses from the base to the 10.3 m tree level height are around 20–22 growth rings and 42±2 mm. At the top of the tree, the sapwood thickness is narrower. The heartwood, which decreases in thickness with increasing tree level heights is not found at the top of the tree. The heartwood appears as a conical shape in the tree trunk. There is no statistical difference in sapwood/heartwood thickness between E-W aspects. Tree level heights and the tree level age were found to be important parameters in determining the thickness of sapwood/heartwood.  相似文献   

3.
  • The wood economics spectrum provides a general framework for interspecific trait–trait coordination across wide environmental gradients. Whether global patterns are mirrored within species constitutes a poorly explored subject. In this study, I first determined whether wood density co‐varies together with elevation, tree growth and height at the within‐species level. Second, I determined the variation of wood density in different stem parts (trunk, branch and twigs).
  • In situ trunk sapwood, trunk heartwood, branch and twig densities, in addition to stem growth rates and tree height were determined in adult trees of Nothofagus pumilio at four elevations in five locations spanning 18° of latitude. Mixed effects models were fitted to test relationships among variables.
  • The variation in wood density reported in this study was narrow (ca. 0.4–0.6 g cm?3) relative to global density variation (ca. 0.3–1.0 g cm?3). There was no significant relationship between stem growth rates and wood density. Furthermore, the elevation gradient did not alter the wood density of any stem part. Trunk sapwood density was negatively related to tree height. Twig density was higher than branch and trunk densities. Trunk heartwood density was always significantly higher than sapwood density.
  • Negative across‐species trends found in the growth–wood density relationship may not emerge as the aggregate of parallel intraspecific patterns. Actually, trees with contrasting growth rates show similar wood density values. Tree height, which is tightly related to elevation, showed a negative relationship with sapwood density.
  相似文献   

4.
The susceptibility of Scots pine (Pinus sylvestris L.) sap- and heartwood against the wood decaying brown-rot fungus (Coniophora puteana) was investigated after long-term forest fertilization at three different sites in central Finland. Different wood properties: wood extractives, wood chemistry, and wood anatomy were used to explain sap- and heartwood decay. Scots pine sapwood was more susceptible to decay than its heartwood. In one site, sapwood seemed to be more resistant to wood decay after forest fertilization whereas the susceptibility of heartwood increased. Significant changes in the sapwood chemistry were found between treatment and sites, however, no relationship between wood chemistry and wood decay was observed in the factor analysis. The results of this study show that there was an inconsistent relationship between decay susceptibility and fertilization and the measured physical and chemical attributes of the wood were not consistently correlated with the decay rate.  相似文献   

5.
《Annals of botany》1997,79(5):479-485
The purpose of this paper is to determine the extent to which the physical and mechanical properties of dry and green wood samples are correlated. Samples of green (fresh) sap- and heartwood differing in density (ρ) were removed from the trunk of a black locust (Robinia pseudoacaciaL.) tree 30 years old and measuring 15 m in height. These samples were mechanically tested to determine their Young's elastic modulus (E), proportional (elastic) limit (σp), and modulus of rupture (σR). The Young's elastic modulus of green wood samples increased in magnitude to a limit with increasing cross-sectional area of the sample tested. The values of all three mechanical parameters measured for sapwood samples were consistently lower than those measured for heartwood samples with equivalent cross-sectional areas.Ewas linearly and positively correlated with the σpand σRof heartwood tissue samples. All mechanical properties were highly correlated with the density of green heartwood. Likewise, these properties were highly correlated with the specific gravity of wood samples. Based on these results, it is concluded that either the density of fresh wood or the specific gravity of air-dried wood can be used to estimate the mechanical properties of black locust wood based on simple regression curves in the absence of extensive mechanical tests.  相似文献   

6.
Nitrogen, phosphorus, potassium, calcium, and magnesium concentrations in woody tissue are poorly documented, but are necessary for understanding whole-tree nutrient use and storage. Here, we report how wood macronutrient concentrations vary radially and along the length of a tree for 10 tropical tree species in Sabah, Malaysia. Bark nutrient concentrations were consistently high: 2.9–13.7 times greater than heartwood depending on the nutrient. In contrast, within the wood both the radial (sapwood vs. heartwood) and vertical (trunk bottom vs. trunk middle) variation was modest. Higher concentrations in sapwood relative to heartwood provide empirical support for wood nutrient resorption during sapwood senescence. Dipterocarp species showed resorption rates of 25.3 ± 7.1% (nitrogen), 62.7 ± 11.9% (phosphorus), and 56.2 ± 12.5% (potassium), respectively, while non-dipterocarp species showed no evidence of nutrient resorption in wood. This suggests that while dipterocarps have lower wood nutrient concentrations, this family is able to compensate for this by using wood nutrient resorption as an efficient nutrient conservation mechanism. In contrast to other nutrients, calcium and magnesium tended to accumulate in heartwood. Wood density (WD) showed little vertical variation along the trunk. Across the species (WD range of 0.33 to 0.94 mg/cm3), WD was negatively correlated with wood P and K concentration and positively correlated with wood Ca concentration. As our study showed exceptionally high nutrient concentrations in the bark, debarking and leaving the bark of the harvested trees on site during logging operations could substantially contribute to maintaining nutrients within forest ecosystems.  相似文献   

7.
Hydraulic specific conductivity, vulnerability to cavitation and water storage capacity of Douglas-fir sapwood was determined for samples from six young (1.0-1.5 m tall) and six mature trees (41-45 m tall). Measurements on samples from young trees showedthere were no effects of two contrasting sample types (entire stem segments vs sectors chiseled out of entire stems) on any of the calculated hydraulic parameters, for vulnerability to cavitation and water storage capacity. Measurements on mature trees were made on wood from four heights on the bole and from two sapwood depths. Outer and inner sapwood at the base of the tree had higher water storage capacities and were more vulnerable to cavitation than was sapwood from the tree top. At every height, old trees were more vulnerable to cavitation than at 1.0 m from the ground in young trees. The water storage capacities showed three distinct phases at the base of the trunk. Young trees had similar water storage capacity (per unit volume of sapwood) to the topof the mature trees, which was lower than the water storage capacity throughout the rest of the bole xylem. The way in which capacitance was calculated (on a volumetric basis vs a relative water content basis) affected the conclusion one would draw at the low water potentials (<-3 MPa). Within a tree, we found an apparent trade-off between having both hydraulic specific conductivity and stem water storage, and vulnerability to cavitation.  相似文献   

8.
Wood is of critical importance to humans as a primary feedstock for biofuel, fiber, solid wood products, and various natural compounds including pharmaceuticals. The trunk wood of most tree species has two distinctly different regions: sapwood and heartwood. In addition to the major constituents, wood contains extraneous chemicals that can be removed by extraction with various solvents. The composition and the content of the extractives vary depending on such factors as, species, growth conditions, and time of year when the tree is cut. Despite the great commercial and keen scientific interest, little is known about the tree-specific biology of the formation of heartwood and its extractives. In order to gain insight on the molecular regulations of heartwood and its extractive formation, we carried out global examination of gene expression profiles across the trunk wood of black locust (Robinia pseudoacacia L.) trees. Of the 2,915 expressed sequenced tags (ESTs) that were generated and analyzed in the current study, 55.3% showed no match to known sequences. Cluster analysis of the ESTs identified a total of 2278 unigene sets, which were used to construct cDNA microarrays. Microarray hybridization analyses were then performed to survey the changes in gene expression profiles of trunk wood. The gene expression profiles of wood formation differ according to the region of trunk wood sampled, with highly expressed genes defining the metabolic and physiological processes characteristic of each region. For example, the gene encoding sugar transport had the highest expression in the sapwood, while the structural genes for flavonoid biosynthesis were up-regulated in the sapwood-heartwood transition zone. This analysis also established the expression patterns of 341 previously unknown genes.  相似文献   

9.
The amounts of free sterols, steryl esters and lipid phosphorus were determined in the sapwood and heartwood of mature, and in the outer and inner sapwood of young Pinus sylvestris trees. In the mature trees (up to 70 years old) the heartwood contains significantly higher amounts of free sterols than the sapwood. No radial gradient can be demonstrated in the amounts of steryl esters. Lipids extracted from the sapwood contain higher amounts of phosphorus than those from the heartwood. Stems of young Pinus sylvestris trees (up to 13 years old) show in the inner sapwood higher amounts of both free sterols and steryl esters than the peripheral younger wood zone. The inner sapwood of the young stems shows slightly higher amounts of lipid phosphorus than the outer sapwood. The results indicate that Pinus sylvestris accumulates both free sterols and steryl esters in the stems at a very early stage of the life cycle. Sterol accumulation in the innermost parts of the stems seems not to depend on heartwood formation.  相似文献   

10.
Summary The distributions of reserve carbohydrates and of three dominant heartwood extractives were determined in the trunkwood of Robinia pseudoacacia L. The trees were cut at different times of the year (September, November, January, and April). With the exception of the tree felled in January, all trunks exhibited highest contents of nonstructural storage carbohydrates (glucose, fructose, sucrose, and starch) in the youngest, outermost sapwood zone. With increasing depth of the trunk, the levels of carbohydrates decreased. At the sapwood-heartwood transition zone, only trace amounts of nonstructural carbohydrates were present. The heartwood itself contained no storage material. The wood zones of different ages of the trees cut in September, November, and January exhibited glucose/fructose ratios of approximately 1. In April, however, there was a shift to glucose. In the youngest sapwood the amounts of soluble sugars were higher in the earlythan in the latewood. Older zones of the sapwood and the sap-wood-heartwood transition zone showed the opposite behaviour. Three main wood extractives of Robinia were characterized and quantified: the flavanonol dihydrorobinetin (DHR), the flavonol robinetin (ROB) and a hydroxycinnamic acid derivative (HCA). Only DHR was present — in very low amounts — in the younger sapwood of all trunks investigated. Higher amounts (>1 mol/g dry weight) of this compound and the HCA were present in the sapwood-heartwood transition zone. DHR augmented within the heartwood up to a more or less constant level. HCA increased towards the heartwood and decreased again in the inner heartwood parts. ROB appeared in the innermost parts of the sapwood-heartwood transition zone and reached maximum values in older parts of the heart-wood. The results indicate that starch is hydrolyzed at the sapwood-heartwood boundary and thus represents a primary major source of hydroxycinnamic acid and flavonoid synthesis.Dedicated to Prof. Meinhart H. Zenk on the occasion of his 60th birthday  相似文献   

11.
Eastern redcedar is widespread in the US and produces significant amount of biomass. Open-grown trees invade abandoned fields and compete with valuable forage species in pastures and rangelands. Value-added product development from redcedar is vital for management of eastern redcedar. Cedarwood oil is a valuable component which can be used for further value-added product development. This study examined the effect of age on the distribution of oil in redcedar tree segments. Trunks of eastern redcedar (Juniperus virginiana L.) trees at different stages of growth (26-63 years old) were divided into three sections (top, center and lower). Each section was fractionated separately into bark, heartwood and sapwood segments. Heartwood and sapwood samples from each tree section were analyzed for oil content and composition. A hydrodistillation method was used for oil extraction. Volatile components of tree segments were examined by using a Gas Chromatograph-headspace analysis technique. The heartwood of eastern redcedar contained significantly higher oil than sapwood. Older trees had more oil in the heartwood than younger trees. Both redcedar bark and leaves contained significantly lower oil content than the cedarwood. There were also significant differences in the oil composition of bark, leaves and wood fractions. Cedarwood oil extraction may benefit from prior separation of tree segments prior to oil extraction. However, the economic feasibility of separation prior to an extraction process needs to be further studied. Required extra capital investment and operating costs need to be examined, as well as whether sapwood is worth processing.  相似文献   

12.
The mechanism of heartwood formation in Cryptomeria japonica D. Don has long been studied since heartwood formation is a fundamental physiological feature of trees. In this study, the water distribution in the xylem of C. japonica was investigated at the cellular level to reveal the role of water distribution in the xylem during heartwood formation. Samples were taken from different heights of each trunk, in which the phases of heartwood formation differed. These were designated as SIH, which consisted of sapwood, intermediate wood, and heartwood; SI, which consisted of sapwood and intermediate wood but no heartwood; and S-all, which consisted entirely of sapwood. Cryo-scanning electron microscopic observations of the heartwood-formed (SIH) and non-heartwood-formed (SI and S-all) xylem revealed different patterns of water distribution changes in tracheids between the latewood and earlywood. In the latewood, almost all tracheids were filled with water in all areas from the sapwood to the heartwood (98–100% of tracheids had water in their lumina). In the earlywood, however, the water distribution differed between the sapwood (95–99%), intermediate wood (7–12%), and heartwood (4–100%). Many of the tracheids in the xylem, where the sapwood changed to intermediate wood lost water. In the heartwood, some tracheids remained empty, while others were refilled with water. These results suggest that the water distribution changes in individual tracheids are closely related to heartwood formation. Water loss from tracheids may be an important factor inducing heartwood formation in the xylem of C. japonica.  相似文献   

13.
The distribution of the above-ground and below-ground biomass of Scots pine in southern Finland were investigated in trees of different ages (18–212 years) from two types of growth site. Secondly, some structural regularities were tested for their independence of age and growth site. Trees were sampled from dominant trees which could be expected to have a comparable position in stands of all ages. All stands were on sorted sediments. The biomass of the sample trees (18 trees) was divided into needles, branch sapwood and heartwood, stem sapwood and heartwood, stem bark, stump, large roots (diameter >20 cm), coarse roots (five classes) and fine roots. The amount of sapwood and heartwood was also estimated from the below-ground compartments. Trees on both types of growth site followed the same pattern of development of the relative shares of biomass compartments, although the growth rates were faster on the more fertile site. The relative amount of sapwood peaked after canopy closure, coinciding with the start of considerable heartwood accumulation. The relative amount of needles and fine roots decreased with age. The same was true of branches but to a lesser degree. The relative share of the below-ground section was independent of tree age. Foliage biomass and sapwood cross-sectional area were linearly correlated, but there were differences between the growth sites. Needle biomass was linearly correlated with crown surface area. The fine root to foliage biomass ratio showed an increasing trend with tree age.  相似文献   

14.
A distribution of extractive substances by height on the trunk and radius of the wood of the Siberian larch (Larix sibirica Ledeb.) was investigated. The maximum flavonoid content in terms of dihydroquercetin (the main flavonoid of the larch wood) was shown to be in the butt part of the tree. In the radial direction, this parameter increased from the center to periphery of the heartwood and reached the maximum value at the sapwood boundary. The maximum content of arabinogalactan (AG) was observed in the bottom and in the top of the trunk. The distribution of the extractive substances was studied in roots of the Siberian larch for the first time.  相似文献   

15.
Dendrochronology is a well-established dating method for wooden objects, but due to surface processing of construction timber or natural degradation the dating of historical wood often relies on a prediction of the number of missing rings based on sapwood statistics. Since Scots pine (Pinus sylvestris L.) is one of the most common tree species in north-western Europe, the absence of reliable sapwood statistics and models for the prediction of missing sapwood rings for pine samples is remarkable. We have therefore produced sapwood statistics based on data from 776 pine trees with ages from 15 to 345 years. The material consists of both living trees and historical timber, with varying growth rates, geographic settings, and from different soil types. When the whole material is considered, the average age of the trees is 103 years, and the number of sapwood rings is 54 ± 15 (1 SD), but range from 18 to 129. Trees less than 100-years in age contained 46 ± 11 (1 SD) sapwood rings and had an average tree-ring width (TRW) of 1.76 mm. With increasing age, the average TRW decreased while the number of sapwood rings increased. The average TRW of 101–200-year-old trees is 0.99 mm while the samples contained 63 ± 12 (1 SD) sapwood rings. For trees older than 201 years, the average TRW is 0.64 mm while the number of sapwood rings increased to 85 ± 16 (1 SD). The two most important factors in determining the number of sapwood rings for a given tree when only heartwood statistics are available proved to be (i) the number of heartwood rings and (ii) the average TRW of the heartwood rings. For incomplete samples, we have therefore developed a statistical model based on the sample’s heartwood rings (number and average width) to compute a prediction interval for the total number of rings. The sapwood and heartwood statistics suggest a statistical model for the number of sapwood rings with mean that increase with the number of heartwood rings. Furthermore, the average number of sapwood rings decreases with the mean width of the heartwood rings. However, the predictive power of the mean width is limited when the number of heartwood rings has already been taken into account. Thus, we suggest making predictions for the number of sapwood rings using only the number of heartwood rings. Predictions of the number of sapwood rings based on the statistical model where convincing in the case of the three different datasets that were analysed. The certainty in these predictions was such that the width of the 80% and 95% prediction intervals ranged 28–34 and 45–52 sapwood rings, respectively. Additionally, we demonstrate how make predictions when there is information about the number of remaining sapwood rings in a given sample. To make the sapwood model available, we present a free online R package for fitting our models and an online software dashboard.  相似文献   

16.
Ponderosa pine has very wide sapwood, and yet the spatial and temporal use of that sapwood for water transport is poorly understood. Moreover, there have been few comparisons of function in tips of old-growth trees in comparison with young trees. In the present study, axial and radial specific conductivity (ks), leaf specific conductivity (LSC), leaf specific conductance (kl), native embolism and the compartmentalization of sapwood water storage were characterized in trunks of young and old-growth trees. Trunks of young trees had lower ks, lower LSC and lower native embolism [corresponding to 5% loss of conductivity (PLC)] than trunks of old-growth trees. However, kl in young trees was 3.5 times higher than in old-growth trees, supporting the hypothesis that tall trees have a reduced ability to transport water to their leaves. Water storage (capacitance) of young trees was not significantly different than at the base of old-growth trees. Although the top of the old-growth trees had similar ks, LSC and kl to the young trees for a given cambial age, they had higher native embolism and lower capacitance. There was no trade-off between ks and native embolism at any height. In the tree crown, outer sapwood had 35–50% higher ks than the inner sapwood and 17–25 PLC lower native embolism. At the base of the old trees, there was no significant difference in native embolism between the outer, middle and inner sapwood, showing that refilling of embolisms was complete despite the 130-year difference in wood age among these radial positions. Although during the dry season the inner sapwood tended to be more saturated than the outer sapwood, the outer part of the sapwood contributed up to 60% of the overall stored water. Safer xylem, higher capacitance and higher kl would appear adaptive in the young trees for regulating their water resource, which is likely to be less reliable than the water availability of older trees with their more developed root system.  相似文献   

17.
基于黑龙江省林口林业局、东京城林业局和东北林业大学帽儿山实验林场的35株人工长白落叶松的解析样木数据,构建长白落叶松的边材、心材和树皮密度的Beta回归模型,采用赤池信息准则、决定系数、平均绝对偏差、均方根误差和似然比检验对模型的拟合优度进行比较评价,进而选取边材、心材和树皮密度的最优模型,最后采用刀切法对选择出的最优模型进行检验,评价模型预测能力。结果表明: 边材、心材和树皮密度的最优模型的自变量不完全相同,其中,边材密度与树木年龄、树高、相对高度和相对高度的平方关系较好,而心材密度最优模型的自变量为年生长量、相对高度和相对高度的平方,树皮密度最优模型的自变量为树木年龄、年生长量、相对高度和相对高度的平方。对最优模型分析可知,从树干基部到树梢,边材密度逐渐减小,心材密度先减小后增加,树皮密度先增加后减小。本研究所建立的Beta回归模型可以预估该研究区域的人工林内长白落叶松的边材、心材和树皮任意位置的木材密度,为树干平均密度和生物量的研究奠定基础。  相似文献   

18.
Elucidation of the mechanisms involved in long-distance water transport in trees requires knowledge of the water distribution within the sapwood and heartwood of the stem as well as of the earlywood and latewood of an annual ring. X-ray computed tomography is a powerful tool for measuring density distributions and water contents in the xylem with high spatial resolution. Ten- to 20-year-old spruce (Picea abies L. KARST.) and oak (Quercus robur) trees grown in the field were used throughout the experiments. Stem and branch discs were collected from different tree heights, immediately deep frozen, and used for the tomographic determinations of spatial water distributions. Results are presented for single-tree individuals, demonstrating heartwood and sapwood distribution throughout their entire length as well as the water relations in single annual rings of both types of wood. Tree rings of the sapwood show steep water gradients from latewood to earlywood, whereas those of the heartwood reflect water deficiency in both species. Although only the latest two annual rings of the ringporous species are generally assumed to transport water, we found similar amounts of water and no tyloses in all rings of the oak sapwood, which indicates that at least water storage is important in the whole sapwood.  相似文献   

19.
When lodgepole pines (Pinus contorta Douglas ex Louden var. latifolia Engelm. ex S. Watson) that are killed by the mountain pine beetle (Dendroctonus ponderosae) and its fungal associates are not harvested, fungal decay can affect wood and fibre properties. Ophiostomatoids stain sapwood but do not affect the structural properties of wood. In contrast, white or brown decay basidiomycetes degrade wood. We isolated both staining and decay fungi from 300 lodgepole pine trees killed by mountain pine beetle at green, red, and grey stages at 10 sites across British Columbia. We retained 224 basidiomycete isolates that we classified into 34 species using morphological and physiological characteristics and rDNA large subunit sequences. The number of basidiomycete species varied from 4 to 14 species per site. We assessed the ability of these fungi to degrade both pine sapwood and heartwood using the soil jar decay test. The highest wood mass losses for both sapwood and heartwood were measured for the brown rot species Fomitopsis pinicola and the white rot Metulodontia and Ganoderma species. The sap rot species Trichaptum abietinum was more damaging for sapwood than for heartwood. A number of species caused more than 50% wood mass losses after 12 weeks at room temperature, suggesting that beetle-killed trees can rapidly lose market value due to degradation of wood structural components.  相似文献   

20.
Heat pulse velocity (HPV) apparatus was used to investigatethe relation between the radial HPV profile and the positionof growth rings in two Pinus patula trees (aged 4 and 7 years),growing in Frankfort State Forest in the eastern Transvaal,South Africa. Successive measurements of HPV at 3.0 mm intervalsalong each of four radii in each tree revealed a cyclical pattern,with maxima and minima correlated with summer and winter wood,respectively. We recommend that probe emplacement in fast-growingpines with relatively wide growth rings should be to randomly-assigneddepths in the sapwood. Confidence limits associated with samplemeans of wound-corrected HPV obtained with 4, 8 or 12 probesets were generated by simulating the repeated selection ofa randomly-chosen HPV measurement from each sapwood radius.We conclude from this analysis that a minimum of 12 point measurementsper tree is required to obtain an acceptable estimate of sapflow. In older trees where a decline in HPV is expected towardsthe centre of the trunk, sampling must be modified to avoidbias caused by deeper measurements representing proportionatelyless of the total sapwood area than those taken at more shallowdepths in the sapwood. Key words: Pinus patula, heat pulse velocity, sap flow, growth rings  相似文献   

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