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1.
West, Brown and Enquist (1999a) modeled vascular plants as a continuously branching hierarchical network of connected links (basic structural units) that ends in a terminal unit, the leaf petiole, at the highest link order (WBE model). We applied the WBE model to study architecture and scaling between links of the water transport system from lateral roots to leafy lateral branches and petioles in Populus deltoides Bartr. ex Marsh. trees growing in an agroforestry system (open-grown trees) and in a dense plantation (stand-grown trees). The architecture of P. deltoides violates two WBE model assumptions: (1) the radii of links formed in a branching point are unequal; and (2) there is no terminal unit situated at the end of a hierarchical network, rather, petioles are situated at any link order greater than 1. Link cross sections were taken at various link orders and morphological levels in roots and shoots of open-grown trees and shoots of stand-grown trees. Scaling of link radii was area-preserving. From roots to branches, vessel diameters were scaled with link order in accordance with a 1/6-power, as predicted by the WBE model indicating general vessel tapering. However, analysis of the data at the morphological level showed that vessel radius decreased intermittently with morphological level rather than continuously between successive link orders. Estimation of total water conductive area in a link is based on conducting area and petiole radius in the WBE model. The estimation failed in P. deltoides, probably because petioles are not a terminal unit. Biomass of stand-grown trees scaled with stem basal radius according to the 3/8-power predicted by the WBE model. Thus, the WBE model adequately described vascular allometry and biomass at the whole-tree level in P. deltoides despite violation of Assumption 1, but failed in predictions where the leaf petiole was used as a terminal unit.  相似文献   

2.
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   

3.
In plants, water flows from roots to leaves through a complex network of xylem conduits. The xylem architecture is characterized by the conduit enlargement towards the stem base and the multiplication of conduits near the apices of lateral branches. The xylem architecture of a small ash tree was analysed by measuring the vessel hydraulic diameter (Dh) and number (N) at different heights along the stem and branches. Along the stem, Dh and N increased from the apex to the point of crown insertion. Below, Dh and N decreased and remained constant, respectively. In branches, the Dh and N of apices increased with distance from the ground (PL) (P < 0.001 and P < 0.0001, respectively), indicating that apical resistance (R(APEX)) becomes lower in the most peripheral branches (P < 0.0001). At the level of branch nodes along the stem, the total conductive area (AC) of the stem and branches just above the node was 11% higher than that of the stem just below the node (P = 0.024), whereas the conductivity (Kh) remained invariant above and below (P = 0.76). The difference in AC (ΔAC) between the branches and stem above each node increased with the distance of the node position from the stem apex (L). The xylem architecture of the analysed tree was characterized by anatomical modifications likely aimed at equilibrating the different path length effects on the hydraulic resistance of the different branches. Conduit tapering and multiplication seem to play a crucial role for the achievement of equal hydraulic resistance of all the leaves in the crown.  相似文献   

4.
Phenotypes more or less deficient in photosynthetic pigments show reduced productivity. Not much is known, however, about the influence of pigment-less twigs on the water balance of whole trees. We studied the water relations and hydraulic properties of normal and pigment-less (white) and 1-year-old shoots of 12-year-old Citrus sinensis L. trees. Compared with green leaves, white leaves showed a pronounced deficiency of pigments, higher stomatal density, the absence of chloroplasts in the guard cells and a different organization of leaf parenchyma. Stomatal conductance (gL) and transpiration rate (EL) were higher in white leaves than in green leaves during the hottest hours of the day, especially in July and September. The absence of chloroplasts in the stomatal guard cells seemed to be one of the factors causing insufficient stomatal control. Hydraulic conductance (KL) was higher in white leaves than in green leaves (16.96+/-2.24x10(-5) versus 11.26+/-0.66x10(-5) kg s-1 m-2 MPa). The ratio between the sum of the fourth power of xylem conduit radius (Sigmar4) (which determines theoretical conductance) and the total leaf area supplied by petioles and midribs was higher in white leaves than in green leaves. This was because of a smaller leaf area in white leaves and a statistically different distribution of lumen diameters of the conduits in midribs and petioles. The hydraulic properties of white twigs profoundly disturbed the water balance and physiology of the whole tree.  相似文献   

5.
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   

6.
通过分析初冬园林树木枝条内养分含量变化和落叶内营养元素含量差异,为指导园林树木越冬养护实践和景观设计中科学选择树木种类提供理论依据。选择大庆市小游园5种园林树种,测定枝条养分和落叶养分含量。结果表明:10月26日至11月16日间,丁香枝条氮、磷含量分别降低了68.28%、35.61%;偃伏莱木枝条氮、钾含量分别降低了25.93%、37.40%;垂枝榆枝条氮、磷、钾含量变化表现为先升高后降低,11月较10月降低了24.08%、19.78%、9.32%;榆树枝条磷、钾含量表现为一直升高的变化,升高幅度分别为21.78%、123.86%;落叶中垂枝榆氮含量为1.88%,榆树磷含量为1.60%,榆叶梅钾含量为1.88%。初冬园林树木枝条内养分含量逐渐降低,落叶平衡补充了土壤营养元素。  相似文献   

7.
We developed site-specific allometric models for Leucaena leucocephala × pallida var. KX2 trees in a shaded coffee agroecosystem in Hawaii to predict above- and belowground biomass and the regrowth potential of pollarded trees. Models were used to compare tree growth rates in an experimental agroforestry system with different pollarding frequencies and additions of tree pruning residues as mulch. For all allometric equations, a simple power model (Y = aXb) provided the optimal prediction of biomass or regrowth after pollarding. For aboveground biomass components (stem, branches, leaves, and seed and pods), stem diameter alone was the best predictor variable. Stump diameter provided the best prediction of coarse root biomass and aboveground regrowth after pollarding. Predictions of biomass from generalized allometric models often fell outside the 95% confidence intervals of our site-specific models, especially as biomass increased. The combination of pollarding trees once per year plus the addition of tree mulch resulted in the greatest aboveground regrowth rates as well as accumulation of biomass and C in the stump plus coarse roots. Although optimal prediction required the development of site-specific allometric relationships, a simple power model using stem or stump diameter alone can provide an accurate assessment of above- and belowground tree biomass, as well as regrowth potential under specific management scenarios.  相似文献   

8.
The scaling relationship between leaf area and total mass of plant has important implications for understanding resource allocations in the plant. The model of West, Brown and Enquist(WBE model) considers that a 3/4 scaling exponent of metabolic rate versus total mass to be optimal for each plant and has been confirmed numerous times. Although leaf area is a better proxy of the metabolic rate than leaf mass, few studies have focused on the scaling exponent of leaf area versus total mass and even fewer have discussed the diversification of this scaling exponent across different conditions. Here, I analyzed the scaling exponent of leaf area versus total mass of sample plots across world plants. I found that as the plant grows, it allocates fewer resources to photosynthetic tissues than expected by the WBE model. The results also empirically show that this scaling exponent varies significantly for different plant leaf habit, taxonomic class and geographic region. Therefore, leaf strategy in response to environmental pressure and constraint clearly plays a significant role.  相似文献   

9.
Leaf specific conductivity (LSC; the ratio of stem conductivity (K(P)) to leaf area (A(L))), a measure of the hydraulic capacity of the stem to supply leaves with water, varies with soil water content. Empirical evidence for LSC responses to drought is ambiguous, because previously published results were subject to many confounding factors. We tested how LSC of similar-sized trees of the same population, under similar climatic conditions, responds to persistently wet or dry soil. Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) trees were compared between a dry site and a wet site in the Valais, an inner alpine valley in Switzerland. Soil water strongly influenced A(L) and K(P) and the plant components affecting K(P), such as conduit radius, conduit density and functional sapwood area. Trees at the dry site had lower LSC than trees with the same stem diameter at the wet site. Low LSC in trees at the dry site was associated with a smaller functional sapwood area and narrower conduits, resulting in a stronger reduction in K(P) than in A(L). These observations support the hypothesis that trees maintain a homeostatic water pressure gradient. An alternative hypothesis is that relatively high investments in leaves compared with sapwood contribute to carbon gain over an entire season by enabling rapid whole-plant photosynthesis during periods of high water availability (e.g., in spring, after rain events and during morning hours when leaf-to-air vapor pressure deficit is small). Dynamic data and a hydraulic plant growth model are needed to test how investments in leaves versus sapwood and roots contribute to transpiration and to maximizing carbon gain throughout entire growth seasons.  相似文献   

10.
Increases in plant size and structural complexity with increasing age have important implications for water flow through trees. Water supply to the crown is influenced by both the cross-sectional area and the permeability of sapwood. It has been hypothesized that hydraulic conductivity within sapwood increases with age. We investigated changes in sapwood permeability (k) and anatomy with tree age and height in the broad-leaved evergreen species Eucalyptus regnans F. Muell. Sapwood was sampled at breast height from trees ranging from 8 to 240 years old, and at three height positions on the main stem of 8-year-old trees. Variation in k was not significant among sampling height positions in young trees. However, k at breast height increased with tree age. This was related to increases in both vessel frequency and vessel diameter, resulting in a greater proportion of sapwood being occupied by vessel lumina. Sapwood hydraulic conductivity (the product of k and sapwood area) also increased with increasing tree age. However, at the stand level, there was a decrease in forest sapwood hydraulic conductivity with increasing stand age, because of a decrease in the number of trees per hectare. Across all ages, there were significant relationships between k and anatomy, with individual anatomical characteristics explaining 33-62% of the variation in k. There was also strong agreement between measured k and permeability predicted by the Hagen-Poiseuille equation. The results support the hypothesis of an increase in sapwood permeability at breast height with age. Further measurements are required to confirm this result at other height positions in older trees. The significance of tree-level changes in sapwood permeability for stand-level water relations is discussed.  相似文献   

11.
Becker P  Gribben RJ  Lim CM 《Tree physiology》2000,20(14):965-967
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   

12.
We investigated the vulnerability of xylem to embolism and the seasonal occurrence of xylem embolism in Italian alder (Alnus cordata Loisel.) by acoustic and hydraulic methods. Wood anatomy was also studied. More than eighty percent of the vessels were less than 50 mm long and no vessels were longer than 120 mm. Mean vessel diameter was 48 micro m. Ultrasound acoustic emissions from root and branch segments dehydrating in air followed a similar pattern: in both tissues, emission peaks were recorded when the relative water content of the xylem was around 0.2. In branches dehydrating in air, xylem embolism increased linearly as water potential decreased. In trees in the field, more than 80 percent of hydraulic conductivity was lost in the tree crowns during winter. Recovery from winter embolism occurred mostly before bud burst. In summer, xylem embolism was low (< 30%) and acoustic emissions from roots, stem and branches of trees in the field were also low.  相似文献   

13.
天然麻栎单木地上生物量模型研究   总被引:2,自引:0,他引:2  
通过对铜陵叶山林场麻栎样木地上生物量调查,以胸径、树高为自变量,地上总生物量、树干、树枝、树叶生物量为因变量,选择相对生长式、幂函数式和多项式为生物量回归模型,拟合各模型参数、相关指数、回归剩余离差,并计算生物量估测误差。结果表明:麻栎树干、树枝、树叶和地上总生物量与胸径、树高存在显著幂函数关系,其方程分别为:树干W=6.571×10-4D1.8473H2.411、树枝W=1.163×10-4D2.9497H1.3223、树叶W=0.0032D1.5148H0.8821、总生物量W=9.354×10-4D2.0825H2.1154。树干与总生物量的预估精度均达90%以上。  相似文献   

14.
Allometric models for dominant shade tree species and coffee plants (Coffea arabica) were developed for coffee agroforestry systems in Matagalpa, Nicaragua. The studied shade tree species were Cordia alliodora, Juglans olanchana, Inga tonduzzi and I. punctata. The models predict aboveground biomass based on diameter at breast height (for trees), and the stem diameter at a height of 15 cm and plant height (for coffee plants). In addition, the specific gravity of the studied species was determined.The total aboveground biomass of the shade trees varied between 3.5 and 386 kg per tree, and between 0.005 and 2.8 kg per plant for coffee. The aboveground biomass components (foliage, branch, and stem) are closely related with diameter at breast height (r > 0.75). The best-fit models for aboveground biomass of the shade trees were logarithmic, with adjusted R 2 between 0.71 and 0.97. In coffee plants, a high correlation was found (r = 0.84) with the stem diameter at 15 cm height, and the best-fit model was logarithmic, as well. The mean specific gravity was 0.52 (± 0.11) for trees and 0.82 (± 0.06) for coffee plants.  相似文献   

15.
We investigated the sapling leaf display in the shade among trees of various leaf lifespans co-occurring under the canopy of a warm-temperate conifer plantation. We measured leaf-area ratio (aLAR) and morphological traits of saplings of evergreen broadleaved tree species and a deciduous tree species. Although we found large interspecific and intraspecific differences in aLAR even among saplings of similar size in the homogeneous light environment, we did not find a consistent trend in aLAR with leaf lifespan among the species. While deciduous trees annually produced a large leaf area, some evergreen broadleaved trees retained their leaves across years and had aLAR values as high as those of deciduous trees. Among leaf-level, shoot-level, and individual-level morphological traits, aLAR was positively correlated with current-year shoots mass per aboveground biomass in deciduous trees, and with the area of old leaves per aboveground mass in evergreen broadleaved trees. Thus, tree-to-tree variation in the degrees of annual shoot production and the accumulation of old leaves were responsible for the interspecific and intraspecific variations in aLAR.  相似文献   

16.
Non-destructive methods for determining the biomass and leaf area of individual trees throughout their growing cycle are an essential tool in agroforestry research, but must be capable of providing reliable estimates despite the influence that management strategies such as pruning may have on tree form. In the present study, allometric methods involving measurements of the diameter of all branches provided reliable estimates of canopy leaf area and biomass for grevillea trees (Grevillea robusta A. Cunn.; Proteaceae) grown as poles, but proved unsuitable for routine measurements because of their time-consuming nature. An alternative, less laborious method based on measurements of trunk cross-sectional area immediately below the first branch of the canopy provided satisfactory allometric estimates of leaf area and canopy biomass. Trunk biomass was determined from measurements of tree height and diameter at breast height using established methodology based on the assumption that trunk volume may be calculated using a quadratic paraboloid model; biomass was determined as the product of trunk volume and the specific gravity of the wood. The theoretical basis, development and validation of allometric methods for estimating tree growth are discussed and their wider applicability to other agroforestry systems is assessed. This revised version was published online in June 2006 with corrections to the Cover Date.  相似文献   

17.
Many authors have attempted to explain the adaptive response of tropical plants to drought based on studies of water relations at the leaf level. Little attention has been given to the role of the xylem system in the control of plant water requirements. To evaluate this role, we studied the hydraulic architecture and water relations parameters of two tropical canopy trees with contrasting leaf phenologies: deciduous Pseudobombax septenatum (Jacq.) Dug and evergreen Ochroma pyramidale (Cav. ex lamb) Urban, both in the family Bombacaceae. The hydraulic architecture parameters studied include hydraulic conductivity, specific conductivity, leaf specific conductivity, and Huber value. Water relations parameters include leaf water potential, stem and leaf water storage capacitance, transpiration, stomatal conductance, and vulnerability of stems to cavitation and loss of hydraulic conductivity by embolisms. Compared to temperate trees, both species showed a pattern of highly vulnerable stems (50% loss of conductivity due to embolism at water potentials less than 1 MPa) with high leaf specific conductivities. The vulnerability of xylem to water-stress-induced embolism was remarkably similar for the two species but the leaf specific conductivity of petioles and leaf-bearing stems of the evergreen species, Ochroma (e.g., 9.08 and 11.4 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively), were 3.4 and 2.3 times higher, respectively, than those of the deciduous species, Pseudobombax (e.g., 2.64 and 5.15 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively). A runaway embolism model was used to test the ability of Ochroma and Pseudobombax stems to maintain elevated transpiration rates during the higher evaporative demand of the dry season. The percent loss of leaf area predicted by the runaway embolism model for stems of Pseudobombax ranged from 5 to 30%, not enough to explain the deciduous phenology of this tree species without analysis of root resistance or leaf and petiole vulnerability to embolism.  相似文献   

18.
Axial water transport in trees is mainly determined by the gradient of negative water pressure and the structure of conductive xylem elements (i.e. conduits) connecting the fine roots with the foliage. There is still an essential lack of knowledge concerning the relationship between wood structure and hydraulic properties, especially of coarse roots. To this end, the study aimed (1) to work out a novel approach, based on the combination of computer tomography (CT) and light microscopy (LM), for determining the cumulative cross-sectional lumen area of conduits involved in the water transport of coarse roots in European beech (Fagus sylvatica) and Norway spruce (Picea abies) and (2) to demonstrate its adequacy in quantifying the functional relationship between sapwood anatomy and ascending water mass flow in the xylem. The cross-sectional sapwood area of coarse roots was assessed through CT. The cumulative cross-sectional lumen area of conduits in the sapwood (i.e. the lumen area of conductive conduits) was measured by LM in combination with interactive image analysis. The new approach was developed with coarse roots of both the tree species growing in a 60-year-old mixed forest in Bavaria, Germany. The combination of the two methods unveiled spruce to possess a distinct sapwood/heartwood boundary in small-diameter roots, whereas such roots of beech reflected a gradual transition zone; only large-diameter roots displayed a distinct boundary in beech. Additionally, the cumulative lumen area of conductive conduits was found to be approximately 12% of the total coarse root cross-sectional area in both the tree species. The new approach of measuring the conductive lumen area of coarse-root conduits yielded levels of specific sap flow (i.e. axial conductivity) that substantially differed from those derived from commonly applied methods, which were based on sap flow per unit of total cross-sectional root area or xylem cross-sectional area of individual roots. The combination of CT and LM will facilitate functional comparisons of woody roots differing in diameter and of tree species of different anatomical xylem structure.  相似文献   

19.
对30种树种在典型大气污染区、重度污染区和清洁区3类不同环境下的叶面积、株高、基径、冠幅等表观指标进行测定分析,结果表明:绝大部分植株生长表观指标均受到大气污染物的影响,对大气污染物敏感性顺序为冠幅〉株高〉基径〉叶面积。植物对大气污染的耐受性有一个阈值。而且得到综合生长比值的计算公式GRC=0.46×RCA+0.37×RH+O.17×RBD。根据此比值和冠幅生长量增减比值,作为划分树种耐受性等级的基准数据,结合植物在典型大气污染区中的生长形态表观(叶、枝、梢、花、果的长势和受害状况)确定了植物对大气污染耐受性等级划分的方法。  相似文献   

20.
Many invertebrates, birds and mammals are dependent on hollow trees. For landscape planning that aims at persistence of species inhabiting hollow trees it is crucial to understand the development of such trees. In this study we constructed an individual-based simulation model to predict diameter distribution and formation of hollows in oak tree populations. Based on tree ring data from individual trees, we estimated the ages when hollow formation commences for pedunculate oak (Quercus robur) in southeast Sweden. At ages of about 200–300 years, 50% of the trees had hollows. Among trees <100 years old, less than 1% had hollows, while all >400-year-old trees had hollows. Hollows formed at earlier ages in fast-growing trees than in slow-growing trees, which may be because hollows are formed when big branches shed, and branches are thicker on fast-growing trees in comparison to slow-growing trees of the same age. The simulation model was evaluated by predicting the frequency of presence of hollows in relation to tree size in seven oak stands in the study area. The evaluation suggested that future studies should focus on tree mortality at different conditions. Tree ring methods on individual trees are useful in studies on development of hollow trees as they allow analysis of the variability in time for hollow formation among trees.  相似文献   

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