Isolation of endophytes from leaves of <Emphasis Type="Italic">Neolitsea sericea</Emphasis> in broadleaf and conifer stands

Endophytic fungi were isolated from leaves of Neolitsea sericea, a major lauraceous tree in the laurel forests of southern Kyushu, collected from the understory layer of broadleaf and conifer stands. Cytosphaera sp. and a species of Ascomycetes in leaf blade segments, plus a xylariaceous species and Phomopsis spp. in petiole segments, were isolated at relatively high frequency. In general, isolation frequencies of endophytes were higher in petiole than blade segments. In blade segments, patterns of endophyte isolation were quite different among stands, while relatively similar in petiole segments. Significant effects of sampling sites or canopy vegetation were rarely detected. the understory layer of laurel forests. Neolitsea sericea is an evergreen broadleaf lauraceous tree, widely distributed in areas of eastern Asia. In southern Kyushu, it is one of the most common trees, growing both as canopy and understory species. In this study, endophytes were isolated from the leaves of N. sericea growing in the understory layer of conifer and broadleaf forest stands to survey the endophytes of N. sericea leaves and to examine the effect of the canopy layer on endophytic mycobiota in understory plants.     

Sun-shade adaptability of the red mangrove,Rhizophora mangle (Rhizophoraceae): Changes through ontogeny at several levels of biological organization

Rhizophora mangle L., the predominant neotropical mangrove species, occupies a gradient from low intertidal swamp margins with high insolation, to shaded sites at highest high water. Across a light gradient, R. mangle shows properties of both “light-demanding” and “shade-tolerant” species, and defies designation according to existing successional paradigms for rain forest trees. The mode and magnitude of its adaptability to light also change through ontogeny as it grows into the canopy. We characterized and compared phenotypic flexibility of R. mangle seedlings, saplings, and tree modules across changing light environments, from the level of leaf anatomy and photosynthesis, through stem and whole-plant architecture. We also examined growth and mortality differences among sun and shade populations of seedlings over 3 yr. Sun and shade seedling populations diverged in terms of four of six leaf anatomy traits (relative thickness of tissue layers and stomatal density), as well as leaf size and shape, specific leaf area (SLA), leaf internode distances, disparity in blade–petiole angles, canopy spread: height ratios, standing leaf numbers, summer (July) photosynthetic light curve shapes, and growth rates. Saplings showed significant sun/shade differences in fewer characters: leaf thickness, SLA, leaf overlap, disparity in bladepetiole angles, standing leaf numbers, stem volume and branching angle (first-order branches only), and summer photosynthesis. In trees, leaf anatomy was insensitive to light environment, but leaf length, width, and SLA, disparities in bladepetiole angles, and summer maximal photosynthetic rates varied among sun and shade leaf populations. Seedling and sapling photosynthetic rates were significantly depressed in winter (December), while photosynthetic rates in tree leaves did not differ in winter and summer. Seasonal and ontogenetic changes in response to light environment are apparent at several levels of biological organization in R. mangle, within constraints of its architectural baiiplan. Such variation has implications for models of stand carbon gain, and suggest that response flexibility may change with plant age.     

Primary ecological succession in vascular epiphytes: The species accumulation model

Epiphytes are integral to tropical forests yet little is understood about how succession proceeds in these communities. As trees increase in size they create microhabitats for late‐colonizing species in both small and large branches while maintaining small tree microhabitats for early colonizing species in the small and young branches. Thus, epiphyte succession may follow different models depending on the scale: at the scale of the entire tree, epiphytes may follow a species accumulation model where species are continuously added to the tree as trees increase in size but at the scale of one zone on a branch (e.g., inner crown: 0–2 m from the trunk), they may follow the replacement model of succession seen in terrestrial ecosystems. Assuming tree size as an indicator of tree age, I surveyed 61 Virola koschnyi trees of varying size (2.5–103.3 cm diameter at breast height) in lowland wet tropical forest in Costa Rica to examine how epiphyte communities change through succession. Epiphyte communities in small trees were nested subsets of those in large trees and epiphyte communities became more similar to the largest trees as trees increased in size. Furthermore, epiphyte species in small trees were replaced by mid‐ and late‐successional species in the oldest parts of the tree crown but dispersed toward the younger branches as trees increased in size. Thus, epiphyte succession followed a replacement model in particular zones within treecrowns but a species accumulation model at the scale of the entire tree crown.     

常绿阔叶林植物叶片N、P化学计量特征对毛竹扩张的响应

为从生态化学计量内平衡角度解释常绿阔叶林不同层次植物对毛竹(Phyllostachys edulis)扩张的生存响应差异性,该研究采用空间代替时间的方法,在江西井冈山国家级自然保护区沿毛竹扩张方向选取典型毛竹-常绿阔叶林界面,依次设置毛竹林、竹阔混交林和常绿阔叶林样地,比较分析了毛竹扩张方向上样地内不同乔木层、灌木层、草本层植物叶片及土壤N、P含量及比例。结果表明:(1)从毛竹林到阔叶林,土壤N含量上升,P含量下降,N:P上升(P<0.05); 乔木层树种 [红楠(Machilus thunbergii)、赤杨叶(Alniphyllum fortunei)及交让木(Daphniphyllum macropodum)]叶片P含量下降,N:P上升(P<0.05); 除灌木层的红果山胡椒(Lindera erythrocarpa)外,各林分中的灌木层和草本层植物N、P含量及比例变化较小。(2)土壤N:P与乔木层、草本层和灌木层植物叶片N:P分布呈显著正相关、负相关与不相关。(3)在各林分中,毛竹叶片N、P含量及比例较稳定。综上认为,毛竹通过改变土壤N、P化学计量特征进行扩张,引起植物体N、P元素化学计量特征发生变化。灌木及草本植物受土壤异质性影响较小,但是乔木层植物N、P元素化学计量特征却因此失衡,这可能是阔叶林乔木层树种存亡受威胁的重要原因。     

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.     

Drought alters the canopy architecture and micro-climate of Hevea brasiliensis trees

 In this study a comparison of the canopy architecture and the growth and distribution of roots was made in 10-year-old trees of Hevea brasiliensis grown in a severely drought-prone area on the west coast of India under rainfed and irrigated conditions. LAI and light interception increased significantly in the irrigated compared to the rainfed trees. Girth and height of the tree were 29 and 19% more while width and height of the canopy were 19 and 20% more in the irrigated than rainfed trees. There were 22% more primary branches which had 26% more diameter in the irrigated trees than rainfed trees. The branches were inserted on the main trunk at an angle of 58.36° in the irrigated and 44.22° in rainfed trees. The above changes led to more light penetration which altered the light distribution inside the rainfed trees during summer and inhibited leaf photosynthesis particularly in the top canopy leaves. In the rainfed trees most of the growth occurred during the short favorable season immediately after the monsoon between June and October and no growth or even shrinking of the trunk was seen during summer. In the irrigated trees a higher growth was seen throughout the year and summer had no adverse effect. Although there was some difference in the root distribution pattern, the total root density per unit soil volume did not vary between the irrigated and rainfed trees. Key words  Hevea brasiliensis· Drought · Crown architecture · Micro-climate · Root growth Received: 8 May 1998 / Accepted 8 October 1998     

Foliar growth of Eriocnema fulva Naudin (Melastomataceae) in a forest fragment in southeastern Brazil.

Eriocnema fulva Naudin is an endangered herbaceous, perennial, iteroparous, evergreen species geographically restricted to southeastern-center Minas Gerais State, SE Brazil. The individuals occur as patches on rocky riverbanks shaded by seasonal semideciduous Atlantic forest; they are fixed by roots and have a pending stem. Aiming to investigate leaf development and its importance for individual survival, fifteen contiguous plots (1 x 1 m) were set down in Jambreiro Forest (19 degrees 58'-59' S and 43 degrees 52'-55' W, 800-1100 m altitude), in the municipality of Nova Lima. A total of 260 individuals with the largest leaf blade length > or = 1 cm was tagged and measured in 1997, 1998, and 1999. Leaf expansion was recorded each month during 26 months until April 2000. Plant size was measured through leaf blade length, petiole length, stem length, and number of leaves. Significant changes were detected only after two years, thus indicating that plant growth is slow. The proportion of surviving leaves after two years was 60%. Total blade expansion took over 14 months, a slow growth rate when compared to leaves of other tropical forest canopy and understory species. Long leaf lifespans are to be found in plants exhibiting slow growth, and we observed that some leaves lived longer than three years. Petiole growth can help to better position the leaf in the search for light, thus contributing to the growth and survival of the plant. The relationships among size measures were significant, reinforcing the great contribution of leaf size for plant size. The age of the largest individual was estimated as 36 years based on the median annual leaf production rate.     

Changes in crown development patterns and current-year shoot structure with light environment and tree height in Fagus crenata (Fagaceae)

The relative effects of light and tree height on the architecture of leader crowns (i.e., the leading section of the main trunk, 100 cm in length) and current-year shoots for a canopy species, Fagus crenata, occupying both the ridge top and the valley bottom in a cool-temperate forest in Japan were investigated. For leader crowns, the number of current-year shoots and leaves increased with increasing tree height, whereas the mean length of current-year shoots increased with increasing relative photon flux density (PFD). The leader crown area decreased, and the depth and leaf area index of leader crowns increased, with increasing relative PFD. The mass of current-year shoots increased with relative PFD. However, this total mass was allocated differently between stems and leaves depending on tree height, such that the relative allocation to stems increased with increasing tree height. Furthermore, stem structures within current-year shoots also changed with height, such that taller trees produced thicker and shorter stems of the same volume. In contrast, leaf structure and leaf biomass allocations changed with relative PFD. Specific leaf area decreased with increasing relative PFD. In addition, leaf number increased more rapidly with increasing individual leaf mass for trees exposed to greater relative PFD. Consequently, the total leaf area supported by a stem of a given diameter decreased with increasing tree height and relative PFD. Thus, the architecture of leader crowns and current-year shoots were related differently to light and tree height, which are considered important for efficient light capture and the growth of small and tall trees in different environments.     

油樟幼苗对马尾松林窗面积的光合响应特征

为了解马尾松人工林窗对伴生树种的影响,为马尾松人工纯林的团块状混交提供科学依据,研究了10 m×10 m(T1)、15 m×15 m(T2)、20 m×20 m(T3)、25 m×25 m(T4)、30 m×30 m(T5)、35 m×35 m(T6)和40 m×40 m(T7)马尾松人工林窗中油樟(Cinnamomum longepaniculatum)幼苗叶片形态和光合生理特征的变化,探讨马尾松林窗斑块对混生树种生长的影响。结果表明:1)林窗面积低于20 m×20 m时,油樟幼苗叶片最大净光合速率显著低于旷地对照;2)叶片比叶重随着林窗面积的增大显著升高;林窗内油樟幼苗叶氮含量在小林窗中(10 m×10 m)显著低于旷地对照,但在大林窗中(如20 m×20 m)显著高于对照;林窗内幼苗叶磷含量则与旷地无显著性差异;3)叶氮在光合组分中的总分配系数随着林窗面积的增加而增大,其中叶氮在羧化组分中的分配系数升高尤为明显,而捕光组分的分配系数在林窗面积10 m×10 m—20 m×20 m范围内随林窗面积的增加而显著降低。可见,当马尾松林窗面积低于20 m×20 m时,林窗环境会显著影响油樟幼苗的光合能力,油樟幼苗可以通过调节比叶面积、叶氮含量以及叶氮在光合组分中的分配等形态、生理适应特征来适应林窗环境的变化。     

Growing tall vs growing wide: tree architecture and allometry of Acacia karroo in forest, savanna, and arid environments

In order to investigate how environmental factors other than light availability affect tree architecture, differences in branching architecture and allometry were analysed in populations of Acacia karroo Hein. from three different environments in South Africa: forests, savannas and arid‐shrublands. Factors such as fire and herbivory have a large effect on tree life history in certain environments and are likely to have selected for trees that have different architectures from those of forest trees, whose major limitation is light assimilation. Significant differences were found in stem architecture and branching architecture between trees in each environment. Compared with forest trees, trees in savannas had an elongated growth form with small canopy and leaf areas, and tall, thin, unbranched trunks. Trees in arid areas showed opposite trends with wider canopies, and increased lateral branching. Savanna trees had significantly smaller spines than trees in other environments, and both forest and savanna trees showed delayed reproduction. These differences are probably related to a trade‐off between an architecture geared towards rapid height‐gain and one promoting lateral spread, and can be explained with reference to the different selective pressures in each environment. In forests, vertical and horizontal growth are both important. However, in savannas there is a great pressure for rapid vertical growth to escape fires, while in arid areas a defensive, lateral growth form is selected for. Savanna trees and arid karoo trees have evolved architectures that are more extreme vertically and laterally than the range of architectures displayed in a forest community.     

Developmental and physiological correlates of leaf size in Hyeronima alchorneoides (Euphorbiaceae)

The tropical emergent tree Hyeronima alchorneoides has large decreases in leaf size with tree age: 1200 cm(2) at 1 yr, 900 cm(2) at 3 yr, 200 cm(2) at 11 yr, and 80 cm(2) in old (>30 yr) individuals. We tracked leaf growth and physiological attributes on trees of three different ages (1, 3, and 11 yr) to determine the developmental basis and functional consequences of this variation. Leaves on young trees grew faster and sustained maximum rates of leaf expansion longer than leaves on older trees. Leaf mass per area (LMA) did not differ among age classes. Maximum photosynthetic rates reflected differences in leaf nitrogen concentration, in which leaves from the lower crown of younger trees outperformed those at a comparable crown position in older trees. One-year-old trees had the lowest stomatal conductance and the greatest instantaneous water use efficiency. Ontogenetic plasticity in mature leaf size, structure, and physiology may be a balance between the advantages conferred by rapid height growth when trees are young and the benefits derived from producing branches that increase light harvesting ability as trees reach the canopy.     

The<Emphasis Type="Italic"> Arabidopsis thaliana rlp</Emphasis> mutations revert the ectopic leaf blade formation conferred by activation tagging of the<Emphasis Type="Italic"> LEP</Emphasis> gene

Activation tagging of the gene LEAFY PETIOLE ( LEP) with a T-DNA construct induces ectopic leaf blade formation in Arabidopsis, which results in a leafy petiole phenotype. In addition, the number of rosette leaves produced prior to the onset of bolting is reduced, and the rate of leaf initiation is retarded by the activation tagged LEP gene. The ectopic leaf blade results from an invasion of the petiole region by the wild-type leaf blade. In order to isolate mutants that are specifically disturbed in the outgrowth of the leaf blade, second site mutagenesis was performed using ethane methanesulphonate (EMS) on a transgenic line that harbours the activation-tagged LEP gene and exhibits the leafy petiole phenotype. A collection of revertant for leafy petiole ( rlp) lines was isolated that form petiolated rosette leaves in the presence of the activated LEP gene, and could be classified into three groups. The class III rlp lines also display altered leaf development in a wild-type (non-transgenic) background, and are probably mutated in genes that affect shoot or leaf development. The rlp lines of classes I and II, which represent the majority of revertants, do not affect leaf blade outgrowth in a wild-type (non-transgenic) background. This indicates that LEP regulates a subset of the genes involved in the process of leaf blade outgrowth, and that genetic and/or functional redundancy in this process compensates for the loss of RLP function during the formation of the wild-type leaf blade. More detailed genetic and morphological analyses were performed on a selection of the rlp lines. Of these, the dominant rlp lines display complete reversion of (1) the leafy petiole phenotype, (2) the reduction in the number of rosette leaves and (3) the slower leaf initiation rate caused by the activation-tagged LEP gene. Therefore, these lines are potentially mutated in genes for interacting partners of LEP or in downstream regulatory genes. In contrast, the recessive rlp lines exhibit a specific reversion of the leafy petiole phenotype. Thus, these lines are most probably mutated in genes specific for the outgrowth of the leaf blade. Further functional analysis of the rlp mutations will contribute to the dissection of the complex pathways underlying leaf blade outgrowth.Communicated by G. Jürgens     

Scaling of Petiole Dimensions with Respect to Leaf Size for a Tropical Tree, Scaphium macropodum (Sterculiaceae), in Borneo

Scaphium macropodum (Miq.) Beumee ex Heyne (Sterculiaceae) in a tropical rain forest in West Kalimantan (Indonesia) was analyzed from the viewpoint of statics. The petiole diameter must increase with increasing leaf size to retain enough mechanical stability and a sufficient amount of conductive vessels. The petiole's cross-sectional area at its base was found to be proportional to the leaf blade's dry mass, which indicates that Shinozaki's pipe model is applicable to leaves with different sizes. Although larger leaves produce greater bending moments on the petiole's cross-section as a result of their greater weights, the bending stresses at the petiole's base caused by the leaf's weight were constant at ca. 76,900 g cm⁻² regardless of leaf size. Thicker petioles increase the leaf's mechanical stability, but require sizable energy investments for their construction. It is hypothesized that the constant value for petiolar stress indicates an optimal balance between energy economy and the mechanical stability of S. macropodum leaves. To keep bending stress constant, the leaf blade's center of gravity shifts to a more proximal position and the cross-sectional area of the petiole increases. Received 8 December 1997/ Accepted in revised form 1 December 1998     

Relationships between branch spacing, growth rate and light in tropical forest saplings

1. The spacing of branches along central stems was related to growth rate and light level in forest saplings and trees in tropical moist forest on Barro Colorado Island (BCI), Panama. The study included 14 species with tiers of plagiotropic branches (having planar leaf arrangements) and four species with continuous distributions of plagiotropic branches.
2. All species showed increases in branch spacing with increasing light and growth rate of diameter, similar to the patterns in leaf spacing noted previously in species which initially bear large leaves on unbranched stems.
3. Non-tiered species had shorter internodes than tiered species but because the latter bear more branches per node, both groups had similar numbers of branches per unit stem length, when compared at similar growth rates.
4. Differences in the relationship between internode length and growth rate among tiered species were related to demographic characteristics, suggesting that tree architecture may influence forest composition.
5. The strong correlation observed between branch spacing and growth rate suggests that branch spacing may be used to estimate past growth histories of forest tree species with plagiotropic branches.     

Influence of leaf size on tree architecture: first branch height and crown dimensions in tropical rain forest trees

 First branch height is an important attribute of sapling architecture, as it defines the height at which prolonged lateral growth is possible. First branch height, measured on saplings of 70 species in tropical rain forests of Australia, Costa Rica, Panama, and Sabah, Malaysia, was highly correlated with leaf blade and petiole length. The observed relationship, first branch height ∝ blade length × (petiole length)^0.5, implies that the ratio of first branch height to blade length increases somewhat with increasing leaf size, among species with a given ratio of petiole to blade length. Orthotropic species, with more or less radially symmetric arrangements of leaves on ascending axes, had a mean first branch height of 7x that observed for plagiotropic species, with planar leaf arrangements. The greater first branch height of orthotropic species was associated with their larger leaves and longer petioles. Plagiotropic species had wider crowns than orthotropic species in the sapling stage, as assessed at the Costa Rican site. Thus, leaf dimensions influence the dynamics of crown construction (or visa versa), as well as affecting leaf energy balance and gas exchange. Received: 5 September 1997 / Accepted: 3 March 1998     

Do holly leaf spines really deter herbivory?

Summary Although spinose teeth of holly leaves have been widely cited as an example of a physical defense against herbivores, this assumption is based largely on circumstantial evidence and on general misinterpretation of a single, earlier experiment. We studied the response of third and fifth instar larvae of the fall webworm, Hyphantria cunea Drury, a generalist, edge-feeding caterpillar, to intact American holly leaves and to leaves that had been modified by blunting the spines, by removing sections of leaf margin between the spines, or by removing the entire leaf margin. The results suggest that the thick glabrous cuticle and tough leaf margin of Ilex opaca are more important than the spinose teeth in deterring edge-feeding caterpillars. Microscopic examination of mature leaves revealed that the epidermis is thickened at the leaf margin, and that the leaf is cirucumscribed by a pair of fibrous veins. In simple choice tests neither domesticated rabbits nor captive whitetailed deer discriminated between spinescent holly foliage and foliage from which spines were removed. Nevertheless, we found little evidence of herbivory by mammals in the field, either on small experimental trees or in the forest understory. While it is possible that spinose teeth contribute to defense by reducing acceptibility of holly relative to other palatable plant species, we suggest that the high concentrations of saponins and poor nutritional quality of holly foliage may be more important than spines in deterring vertebrate herbivores. The degree of leaf spinescence and herbivory was compared at different heights with the tree canopy to test the prediction that lower leaves should be more spinescent as a deterrent to browsers. Leaves on lower branches of mature forest trees were slightly more spinescent than were upper leaves, and juvenile trees were slightly more spinescent than were mature trees. However, there was no relationship between degree of spinescence and feeding damage. The greater spinescence of holly leaves low in the canopy is probably an ontogenetic phenomenon rather than a facultative defense against browsers.The investigation reported in this paper (No. 87-7-8-77) is in connection with a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Dirctor     

LEAF DEVELOPMENT AND CROWN GEOMETRY OF TWO IRIARTEOID PALMS

We examined changes in pinnately compound leaf morphology and crown geometry that occur during height growth of the iriarteoid palms Socratea exorrhiza and Iriartea deltoidea in tropical wet forest of Costa Rica. Although light availability increased with height, the number of leaves per plant was relatively constant. Total leaf area, however, was much larger in taller individuals. Increases in linear dimensions of leaves (length and width) was responsible for less than half of this greater surface area. More important was the transition from a basically dorsiventral display of leaflets in small individuals to a more radial display in taller plants. Production of leaflets in more than one plane resulted in leaves whose surface area was more than twice the horizontally projected area and whose lateral light interception was greatly enhanced. S. exorrhiza, a faster-growing and more light-demanding species, undergoes this transformation in leaf morphology at heights between 3 and 6 m; whereas in the slower-growing and more shade-tolerant I. deltoidea this occurs at heights between 10 and 20 m. Pinnate leaves, with dense radial packing of leaflets along the rachis, are functionally comparable to branches of dicotyledonous trees and may have been important for the evolution of arborescence in palms.     

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。     

Effects of leaf blade narrowness and petiole length on the light capture efficiency of a shoot

Effects of the length: width ratio of a leaf blade and petiole length on shoot light capture were studied with computer simulation. Both a larger length: width ratio and longer petiole contributed to larger light capture per unit leaf area due to a reduced aggregation of leaf area around the stem. Other conditions being equal, shoots with narrow leaves and no petioles and those with wide leaves with petioles showed similar light capture as long as the mean distance of the leaf blade from the stem was the same. In shoots with a short internode and/or distichous phyllotaxis, however, narrow leaves contributed more to avoiding mutual shading than wide leaves with petioles. The predominance of light coming from a higher angular altitude also favored narrow leaves. The possible consequences of these results in the adaptive geometry of plant architecture are discussed.