COMPUTER SIMULATION OF BRANCH INTERACTION AND REGULATION BY UNEQUAL FLOW RATES IN BOTANICAL TREES

Two aspects of branch interaction in trees are investigated theoretically. In the first it is assumed that there is a controlling factor in which the proximity of neighboring terminal branch units influences their branching capability. The almost horizontal tiers of lateral branches of Terminalia catappa L. and Cornus alternifolia L. are simulated by computer using values based on the measured branch geometries of real trees. For branch interaction, we assume a horizontal circle of inhibition whose center is the existing terminal point of a branch. If the end point of another branch extends into the circle, the original branch fails to bifurcate. Examples of computer simulated patterns are illustrated using different degrees of interference and are compared with branch tiers in T. catappa. In the second model the ability of a terminal branch unit to bifurcate is considered to be determined by the accumulation of a critical amount of a hypothetical growth- or bifurcation-determining factor. The daughter branches of a bifurcation are assumed to have differing “flow rates,” i.e., the factor is distributed in different amounts between different daughter axes. Some simulated patterns generated by this model are very similar to real patterns found in T. catappa and an unnamed species of Tabernaemontana. In both simulations bifurcation ratios are determined and are shown to be a variable, not a fixed, property of the simulated trees.     

Leaf variation in a tree of Pourouma tomentosa (Cecropiaceae) in French Guiana

In the flora of French Guiana we find considerable within-plant variation in leaf form. We observed entire, two-lobed, and three-lobed leaves within five separate levels (tiers) of the canopy of a single individual ofPourouma tomentosa subsp.maroniensis. Five branches from each of the five tiers of the tree were collected around the axis of the trunk. From these branches five secondary branchlets were selected and all leaves excised with information recorded as to nodal position, number of leaf nodes, and fertility status of the main branch. This design produced 1015 leaves representing about 20 m² of foliar area and about 2.4 kg of blade dry weight. Our objectives were to determine if statistically significant patterns exist for leaf variation and to suggest improvements for future, general collections. The four lower tiers had 62% entire, 10% 2-lobed, and 28% 3-lobed leaves, in contrast to the top tier with 38% entire, 11% 2-lobed, and 51% 3-lobed leaves. The top tier had no fertile branches. in the lower tiers, fertile branches produced 68% entire leaves whereas nonfertile branches produced only 46% entire leaves. In the top tier, lobed leaves made up 73% of surface area, while in the lower tiers, lobed leaves made up only 48% of total surface area. We selected a random subset of 75 leaves from the 1015, for morphometric analysis using two-way ANOVA (tier×leaf type). The boundaries of leaf images were digitized and rendered into Fourier coefficients, yielding leaf surface area and two variables that quantify aspects of shape: dissection index and leaf complexity. The Fourier coefficients were averaged by tier and by leaf type to reconstruct synthetic, average leaf images. Logistic regression was used to predict the position of leaves on the tree and to provide visualization of the relationships between leaf position on the tree and leaf morphological variables. Within the tree crown, leaf surface area and leaf specific mass (LSM) increases with height, although leaf shape does not change with height. LSM does not vary with leaf form; and sun leaves are larger than shade leaves on this tree. We conducted computer sampling experiments based on exact randomization to simulate the process of obtaining all leaf shapes present in an individual tree when making field collections of varying numbers of duplicates. This also points out the importance of noting the presence of within-tree variation in leaf form on herbarium labeds. Failure to recognize leaf variation can lead to incorrect delimitation of species as well as cause overestimates of the number of species in diversity studies.     

PLASTICITY OF TREE ARCHITECTURE: SPECIFIC AND ECOLOGICAL VARIATIONS FOUND IN AUBREVILLE'S MODEL

The crown form and branching pattern of sapling and adult trees of Terminalia catappa, T. latifolia, and an unidentified species of Manilkara are described. Leaf distribution (leaf positions within the crown) and leaf area index are given for the three species. Although they all exhibit the same architectural model, the two species of Terminalia maintain a tiered canopy in both sapling and large adult trees, while Manilkara has a sapling with poorly defined tiers and an adult with a hollow, hemispherical crown. Thus, the same architectural model (Aubréville's) can give rise to different crown shapes due to quantitative differences in branch geometry, degree of reiteration, and longevity of leaf-bearing branch units. Conversely, similar crown shapes can be produced by trees with different models. Manilkara conforms to the monolayer model of leaf arrangement at maturity, but Terminalia cannot easily be described as a monolayer or a multilayer. Adaptive geometry of a tree and its architectural model are not equivalent nor necessarily comparable.     

Maintenance mechanisms of the pipe model relationship and Leonardo da Vinci’s rule in the branching architecture of <Emphasis Type="Italic">Acer rufinerve</Emphasis> trees

The pipe model relationship (constancy of branch cross-sectional area/leaf area) and Leonardo da Vinci’s rule (equality of total cross-sectional area of the daughter branches and cross-sectional area of their mother branch) are empirical rules of tree branching. Effects of branch manipulation on the pipe model relationships were examined using five Acer rufinerve trees. Half the branches in each tree were untreated (control branches, CBs), and, for the others (manipulated branches, MBs), either light intensity or leaf area (both relating to photosynthetic source activity), or shoot elongation (source + sink activities), was reduced, and responses of the pipe model relationships were followed for 2 years. The pipe model relationship in MBs changed by suppression of source activity, but not by simultaneous suppression of source + sink activities. The manipulations also affected CBs in the year of manipulation and both branches in the next year. The branch diameter growth was most affected by light, followed by shoot elongation and leaf area, in that order. Because of the decussate phyllotaxis of A. rufinerve, one branching node can potentially have one main and two lateral branches. Analysis of 295 branching nodes from 13 untreated trees revealed that the da Vinci’s rule held in branching nodes having one shed branch but not in the nodes without branch shedding, indicating the necessity of natural shedding of branches for da Vinci’s rule to hold. These analyses highlight the importance of the source–sink balance and branch shedding in maintenance of these empirical rules. This article was contributed at the invitation of the Editorial Committee.     

Crown alterations induced by decline: a study of relationships between growth rate and crown morphology in beech (Fagus sylvatica L.)

One of the first symptoms expressed by declining trees is reduced growth in stem diameter and length increment. The possibility of a relationship between length increment and crown thinning in beech (Fagus sylvatica L.) was investigated by developing a computer model to simulate first order branching patterns of the apical 2 m of monopodially branching beech trees, 70–100 years old, for a range of length increment rates. The model was based on values for branching angle, main axis and branch length increment, number of branches produced per year and branch mortality rates for six healthy and declining trees. Shoot growth rates in the apical 2 m of the sample trees ranged from about 5 cm/year (decline class 3) to 43 cm/ year (healthy). Simulations of branching patterns in the apical 2 m of trees growing at different rates indicated that, when growth rate exceeded about 20 cm/year, total first order branch length and area explored were independent of growth rate. When growth rates fell below this value there was a reduction in total area explored and first order branch length due primarily to the formation of fewer branches. More acute branching angles contributed to a reduction in the area explored. Growth rate-related crown thinning could increase the risk of bark necrosis and secondary pathogen infection during dry and/or hot spells.     

The occurrence of non-ring producing branches in Abies lasiocarpa

Lack of annual growth ring production at the base of branches in the lower portion of the crown is a commonly observed phenomenon. In this study, branches with missing rings were found on 47 of 49 Abies lasiocarpa trees sampled. The number of missing rings on the lowest live branch in the crown averaged 12, and ranged as high as 28. The lower one-third of the live crown tended to consist of branches without rings; these branches contained an average of 30% of the total crown leaf area. The ratio of foliage weight to total branch weight, in combination with position of the branch in the canopy, was an effective discriminator of non-ring producing branches. This suggests that both structural and environmental factors influence the cessation of ring production. The potential ecological implications of branches that fail to produce rings are discussed.     

Tree Branching: Leonardo da Vinci's Rule versus Biomechanical Models

This study examined Leonardo da Vinci''s rule (i.e., the sum of the cross-sectional area of all tree branches above a branching point at any height is equal to the cross-sectional area of the trunk or the branch immediately below the branching point) using simulations based on two biomechanical models: the uniform stress and elastic similarity models. Model calculations of the daughter/mother ratio (i.e., the ratio of the total cross-sectional area of the daughter branches to the cross-sectional area of the mother branch at the branching point) showed that both biomechanical models agreed with da Vinci''s rule when the branching angles of daughter branches and the weights of lateral daughter branches were small; however, the models deviated from da Vinci''s rule as the weights and/or the branching angles of lateral daughter branches increased. The calculated values of the two models were largely similar but differed in some ways. Field measurements of Fagus crenata and Abies homolepis also fit this trend, wherein models deviated from da Vinci''s rule with increasing relative weights of lateral daughter branches. However, this deviation was small for a branching pattern in nature, where empirical measurements were taken under realistic measurement conditions; thus, da Vinci''s rule did not critically contradict the biomechanical models in the case of real branching patterns, though the model calculations described the contradiction between da Vinci''s rule and the biomechanical models. The field data for Fagus crenata fit the uniform stress model best, indicating that stress uniformity is the key constraint of branch morphology in Fagus crenata rather than elastic similarity or da Vinci''s rule. On the other hand, mechanical constraints are not necessarily significant in the morphology of Abies homolepis branches, depending on the number of daughter branches. Rather, these branches were often in agreement with da Vinci''s rule.     

ARCHITECTURE AND CROWN GEOMETRY IN TABEBUIA ROSEA (BIGNONIACEAE)

In the tropical tree Tabebuia rosea, seedlings form an erect unbranched stem with rhythmic growth. Three distinct and predictable geometrical stages are then recognized in crown development. Sympodial branching begins with the arrest of the terminal bud of the trunk and symmetric outgrowth of a pair of subtending lateral buds. During an intermediate phase, branching becomes asymmetric at about Order 5. At each sympodial bifurcation there is differentiation between vigorous, relatively straight main branches (leaders) and less vigorous laterals forming regular pseudomonopodial branch complexes, which collectively constitute the cup-shaped crown. Finally, dormant lateral buds in the lowest bifurcation of the trunk are released and reiterate the original crown form. Ultimately an erect, apparently monopodial tree is formed by a set of superimposed cup-shaped crowns. Crown development of Tabebuia is unique because it involves predictable ontogenic changes in branching patterns. Crowns of open-grown Tabebuia consist of relatively few, wide branch tiers, crowns of forest grown trees are tall and narrow. Analysis of the adaptive geometry of wide vs. narrow crowns through computer simulation illustrates the precise cost of mechanical support for terminal leaf rosettes at successive developmental phases and suggests that tall, erect, narrow, and multi-tiered crowns are more efficient than wide open crowns.     

High densities of leaf‐tiers in open habitats are explained by host plant architecture

1. Crown architecture remains one of the least studied plant traits that influence plant–herbivore interactions. The hypotheses that dense crown architecture of mountain birches from open habitats favours leaf‐tying caterpillars through bottom‐up and/or top‐down effects associated with high leaf connectivity were tested. 2. Population densities of leaf‐tying herbivores in open (industrial barren and seashore) habitats were three times as high as in the shaded (forest) habitats. An experimental increase in leaf density by branch binding did not affect foliar consumption by free‐living herbivores but increased consumption by leaf‐tiers. 3. The specific leaf weight was lower in shaded habitats and in bound branches, but branch binding did not influence either the foliar concentrations of carbon and nitrogen or the pupal weight of the most abundant leaf‐tier, Carpatolechia proximella Hbn. (Lepidoptera: Gelechiidae). 4. Caterpillars of C. proximella build several shelters during their lifetime and spend a considerable amount of time outside the shelter, where they excrete most of their faeces. In bound branches, caterpillars built new shelters more frequently than in control branches, and consumed less foliar biomass per shelter. 5. Mortality from parasitoids in bound branches was half that in the control, presumably because the complex environment disrupted parasitoid searching behaviour and/or because of lower damage to leaves from which the shelters were built. 6. It is concluded that the crown architecture associated with high leaf connectivity decreases mortality risks from natural enemies both outside and inside the shelter. Compact and dense crowns of host plants may at least partly explain high population densities of leaf‐tiers in open habitats.     

胡杨枝芽生长特征及其展叶物候特征

以5个不同发育阶段的胡杨(Populus euphratica Oliv.)个体为研究对象,观测记录了枝芽展叶物候、枝芽生长特征和叶形变化的空间分布规律。结果表明:不同发育阶段的胡杨个体以及同一个体树冠的不同层次,其枝芽生长及其展叶物候期表现出不同的时空特征。随着树龄的增加和树冠层次的增高(由基向顶),当年新生枝条长度、枝条叶片数和叶形指数逐渐减小,但叶面积和叶片干重逐渐增大。5个不同发育阶段胡杨个体均表现出展叶物候始于树冠顶层,依次向下结束于树冠基部;展叶物候期共性表现在枝芽萌动期均在4月上旬,起始展叶期集中在4月中旬,展叶终期则在5月上旬到下旬;树龄较大的个体其枝芽萌动期、起始展叶期、展叶终期较树龄较小的个体早;其枝芽萌动期到展叶终期的时间进程较树龄较小的个体短;不同发育阶段的个体枝芽萌动期出现的时间较为离散,起始展叶期和展叶终期出现的时间较为集中。相关分析表明,出叶周期与枝条长度、枝条叶片数量和叶形指数呈极显著正相关,与叶面积和叶片干重呈显著负相关。     

樟子松人工林分枝结构的分析

基于对6块樟子松(Pinus sylvestris var. mongolica)人工林固定标准地中的30株样木枝解析调查数据,通过分析不同林分、不同大小林木1级枝和2级枝的分枝概率、分枝格局和分枝角度,揭示了樟子松人工林树冠的分枝结构特点。研究结果表明：樟子松人工林1级枝和2级枝的平均分枝数量分别为3.84个和2.80个,两者分枝概率均呈正态分布;1级和2级枝条在光照条件好的几个区间（方位角46°～225°）分布较多,1级枝条的水平分布遵从均匀分布,而2级枝条则不遵从均匀分布;树冠上层枝条的分枝角度略小于树冠中、下层,上层平均分枝角度为45.6°,而中下层平均分枝角度都为49.4°。不同大小林木的1级枝分枝结构规律表明：Ⅰ级木和Ⅴ级木的每轮平均分枝数非常接近,分别为3.89和3.94个,比Ⅲ级木每轮分枝数大0.5个左右;1级枝水平分布在各区间内(45°间隔)相差在0.24%～2.81%之间,方差分析结果表明枝条水平分布与林木大小无关;不同大小林木的分枝角度有所差别,Ⅰ级木、Ⅲ级木和Ⅴ级木的平均分枝角度分别为48.5°、42.2°和50.7°。     

Estimation of hydraulic conductance within field-grown apricot using sap flow measurements

Using the heat pulse and other techniques, the hydraulic architecture of apricot trees was mapped out. The flows (overall flow, flow across the four main branches) and forces (water potential differences between xylem and leaves) measured allowed us to quantify hydraulic conductance of branches and of the root/soil resistance. The experiment was carried out in a commercial orchard of 11-year-old apricot trees (Prunus armeniaca L., cv. Búlida, on Real Fino apricot rootstock) during 1 week (October 27–November 3, 1998). Three representative trees with a cylindrical trunk divided into four main branches of different sizes, orientation and local microclimate were chosen for the experiment. Sap flow was measured throughout the experimental period. Twelve sets of heat-pulse probes were used, one for each main branch. The diurnal course of the environmental conditions, the fraction of the area irradiated and leaf water relations were also considered in each main branch. The relationships between leaf water potential, xylem water potential and transpiration were established for different branches and also for the total plant. Using the slopes of these regressions, total plant conductance, the hydraulic conductance of the stem and root pathway, the hydraulic conductance of the canopy and the hydraulic conductance of each branch were estimated. Our findings show that the root conductance and the canopy hydraulic conductance are similar in magnitude. Leaf hydraulic conductance per leaf area unit was similar for each of the four branch orientations, indicating that, while the light microclimate has a dominant influence on transpiration, in this case it had little effect on the hydraulic properties of the canopy.     

Crown structure and vertical foliage distribution in 4-year-old plantation-grown Eucalyptus pilularis and Eucalyptus cloeziana

Tree growth and form are both influenced by crown architecture and how it effects leaf distribution and light interception. This study examined the vertical distribution of foliage in 4-year-old plantation-grown Eucalyptus pilularis Sm. and E. cloeziana F. Muell. trees. Leaf area (LA) distribution was determined at two different sites using allometric approaches to determine LA in crown sections and for whole trees. Leaf area was distributed more towards the upper crowns when canopies had been closed for longer. Leaf area was also skewed more towards the upper crowns for Eucalyptus pilularis than E. cloeziana. These species differences were consistent with differences in vertical light availability gradients as determined by point quantum sensors. Leaf area of individual branches was highly correlated with branch cross-sectional area (CSA) and whole-tree LA was closely related to stem CSA. Branch-level allometric relationships were influenced by site and crown position. However, the general allometric equations between stem size and whole-tree leaf area could be applied across sites. Results from this study suggest that pruning of live branches in these species should follow species-specific guides for the timing and height of pruning to optimise the effects on stem growth and form.     

Architecture and growth of an annual plant <Emphasis Type="Italic">Chenopodium album</Emphasis> in different light climates

Light climates strongly influence plant architecture and mass allocation. Using the metamer concept, we quantitatively described branching architecture and growth of Chenopodium album plants grown solitarily or in a dense stand. Metamer is a unit of plant construction that is composed of an internode and the upper node with a leaf and a subtended axillary bud. The number of metamers on the main-axis stem increased with plant growth, but did not differ between solitary and dense-stand plants. Solitary plants had shorter thicker internodes with branches larger in size and number than the plant in the dense stand. Leaf area on the main stem was not different. Larger leaf area in solitary plants was due to a larger number of leaves on branches. Leaf mass per area (LMA) was higher in solitary plants. It did not significantly differ between the main axis and branches in solitary plants, whereas in the dense stand it was smaller on branches. Dry mass was allocated most to leaves in solitary plants and to stems in the dense stand in vegetative growth. Reproductive allocation was not significantly different. Branch/main stem mass ratio was higher in solitary than dense-stand plants, and leaf/stem mass ratio higher in branches than in the main axis. Nitrogen use efficiency (NUE) (dry mass growth per unit N uptake) was higher and light use efficiency (LUE) (dry mass growth per unit light interception) was lower in the plant grown solitarily than in the dense stand.     

Finding the optimal lengths for three branches at a junction

This paper presents an exact analytical solution to the problem of locating the junction point between three branches so that the sum of the total costs of the branches is minimized. When the cost per unit length of each branch is known the angles between each pair of branches can be deduced following reasoning first introduced to biology by Murray. Assuming the outer ends of each branch are fixed, the location of the junction and the length of each branch are then deduced using plane geometry and trigonometry. The model has applications in determining the optimal cost of a branch or branches at a junction. Comparing the optimal to the actual cost of a junction is a new way to compare cost models for goodness of fit to actual junction geometry. It is an unambiguous measure and is superior to comparing observed and optimal angles between each daughter and the parent branch. We present data for 199 junctions in the pulmonary arteries of two human lungs. For the branches at each junction we calculated the best fitting value of x from the relationship that flow ∞ (radius)^x. We found that the value of x determined whether a junction was best fitted by a surface, volume, drag or power minimization model. While economy of explanation casts doubt that four models operate simultaneously, we found that optimality may still operate, since the angle to the major daughter is less than the angle to the minor daughter. Perhaps optimality combined with a space filling branching pattern governs the branching geometry of the pulmonary artery.     

Spatial variation in sapwood area to leaf area ratio and specific leaf area within a crown of silver birch

Spatial variation in sapwood area to leaf area ratio (Huber value, HV) and specific leaf area (SLA) was examined in branches of closed-canopy trees of silver birch (Betula pendula Roth). HV increased basipetally within a crown and decreased with increasing branch order, but exhibited no significant radial trend along a primary branch. HV was primarily determined by branch position in a crown and branch diameter at the sampling point, being independent of the size of the tree and branch. Greater HV in the lower-crown branches is considered a means to mitigate differences in hydraulic transport capacity between the branches located in different canopy layers. Beside branch position and sampling location on a branch, SLA depended significantly on several other variables characterising tree and branch size. SLA increased basipetally within a crown and along a primary branch, but exhibited no significant trend with branch orders. Because height caused leaf area (A_L) to diminish more rapidly than leaf dry weight, A_L primarily determined the vertical variation in SLA.     

色季拉山急尖长苞冷杉叶片及细根性状随海拔的变异特征

叶片和细根是植物对环境变化响应的主要功能器官,在气候变化趋势下,研究其随环境梯度的变化,对理解植物对环境的适应策略具有重要意义。本文是在色季拉山急尖长苞冷杉分布区,按海拔梯度(3800、3900、4000、4100、4200、4300、4400m)选择7个典型样地,在样地内对其叶片基本特征、叶绿素含量、比叶重和细根特征(0—60cm)等参数进行了测量。结果表明:(1)随海拔梯度升高,急尖长苞冷杉叶片叶面积减小、叶片厚度增加、叶绿素含量逐渐降低、比叶重显著增大。3900m处急尖长苞冷杉的叶片形态特征表现最好,叶片长度、宽度和面积均显著优于其他海拔,海拔4200m叶片厚度达到最大值,叶片面积、叶绿素含量随海拔升高呈下降趋势,但在4200m处出现第二峰值。(2)随海拔梯度增加,细根各性状与海拔表现出非线性关系,其中细根生物量和细根体积在3900m和4200m处出现两次峰值,3900m处细根生物量达到最大值,4200m处细根体积达到最大值,4400m处细根生物量和细根体积均显著少于其他海拔。细根根长密度在海拔3900、4200、4300m较高,三者之间差异不显著,但显著高于其余海拔,4400m海拔细根根长密度最小。细根表面积在3900m海拔处达到最大值,显著高于其他海拔,4200和4300m次之,3800、4000和4400m海拔下细根表面积相对较小。4400m处细根比根长达到最大值。各海拔上细根均主要分布在0—20cm土层。随土层厚度增加,各海拔细根生物量和根系体积在0—60cm土层范围内均逐渐减小;细根根长密度、表面积在20—40cm和40—60cm显著提高;同一海拔细根比根长随土层深度增加呈明显的增加趋势。各海拔40—60cm土层细根比根长显著高于20—40和0—20cm土层。(3)综合叶片及细根特征,海拔3900m为急尖长苞冷杉的最适宜生长区域;随海拔升高,环境因子逐渐恶劣,环境胁迫加剧,急尖长苞冷杉最终形成在4400m处为其分布上限的海拔梯度格局。     

A RELATIONSHIP BETWEEN CIRCUMFERENCE AND WEIGHT IN TREES AND ITS BEARING ON BRANCHING ANGLES

Observation reveals a linear relationship between the logarithm of the circumference of a tree, branch, or leaf stem, and the logarithm of the weight of the tree, branch, or leaf. The bearing of this on the angles of branching in trees is discussed.     

Size structure of current-year shoots in mature crowns

Characteristics of current-year shoot populations were examined for three mature trees of each of three deciduous broad-leaved species. For first-order branches (branches emerging from the vertical trunk) of the trees examined, lengths or diameters of all current-year shoots were measured. Total leaf mass and total current-year stem mass of first-order branches were estimated using an allometric relationship between leaf or stem mass and length or diameter of current-year stems. For each tree, the number of current-year shoots on a first-order branch was proportional to the basal stem cross-sectional area of the branch. On the other hand, first-order branches had shoot populations with size structures similar to each other. As a result, the leaf mass of a first-order branch was proportional to the basal stem cross-sectional area of the branch, being compatible with the pipe-model relationship. All current-year shoot populations had positively skewed size structures. Because small shoots have a larger ratio of leaf mass to stem mass than large shoots, first-order branches had an extremely large ratio of leaf mass to current-year stem mass. This biased mass allocation will reduce costs for current stem production, respiration and future radial growth, and is beneficial to mature trees with a huge accumulation of non- photosynthetic organs. The allometric relationships between leaf mass and basal stem diameter and that between leaf mass and current-year stem mass of first-order branches were each similar across the trees examined. Characteristics of shoot populations tended to offset inter-species diversity of shoot allometry so that branch allometry shows inter-species convergence.     

辽东栎林内不同小生境下幼树植冠构型分析

以黄土高原黄龙山林区辽东栎林内3个小生境（林下、林隙、林缘）下辽东栎天然更新幼树为研究对象,采用典型抽样法对辽东栎幼树侧枝、叶片和树冠的空间分布状况以及生物量分配状况进行调查分析,探讨微生境与幼树植冠构型特征的关系,明确辽东栎幼树对不同小生境的适应策略,为栎林经营和林分结构优化提供理论依据。结果显示：（1）3种生境下辽东栎幼树构型发生了可塑性变化,林下幼树树冠层次比较单一,林隙与林缘的幼树树冠层次更加丰富。（2）由林下至林缘,幼树的树高、枝下高呈逐渐减小的趋势,而地径变化趋势与之相反;幼树的冠幅、树冠面积、树冠率呈先增加后减小的趋势,并且林下与林隙、林缘的差异显著;幼树的总体分枝率、逐步分枝率、枝径比呈先增加后减小的趋势。（3）3种生境下,幼树的一级枝的枝长、直径与倾角随着树高的增加而呈减小的趋势,但3种生境的差异不显著;林下一级枝主要分布在冠层中上部,而林隙与林缘一级枝主要分布在冠层中下、中上部。（4）由林下至林缘幼树叶长、叶宽、单叶面积和比叶面积逐渐降低,而单株叶数、叶总面积、叶面积指数呈先增大后减小趋势;与其他2种生境相比林下叶片分布趋于冠层上部。（5）幼树地上部分生物量中林下主干生物量占83%,枝和叶生物量只占17%;而林隙与林缘虽然各部位生物量有所差异但比例基本一致,其中主干占66%左右,枝和叶生物量占34%左右。研究表明,林隙生境下幼树的构型优于林缘和林下生境,在今后栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化提供有利条件。