Crown plasticity in mixed forests—Quantifying asymmetry as a measure of competition using terrestrial laser scanning

Interspecific competition is a key process determining the dynamics of mixed forest stands and influencing the yield of multispecies tree plantations. Trees can respond to competitive pressure from neighbors by crown plasticity, thereby avoiding competition. We employed a high-resolution ground-based laser scanner to analyze the 3-dimensional extensions and shape of the tree crowns in a near-natural broad-leaved mixed forest in order to quantify the direction and degree of crown asymmetry of 15 trees (Fagus sylvatica, Fraxinus excelsior, Carpinus betulus) in detail. We also scanned the direct neighbors and analyzed the distance of their crown centres and the crown shape with the aim to predict the crown asymmetry of the focal tree from competition-relevant attributes of its neighbors. It was found that the combination of two parameters, one summarizing the size of the neighbor (DBH) and one describing the distance to the neighbor tree (HD), was most suitable for characterizing the strength of the competitive interaction exerted on a target tree by a given neighbor. By summing up the virtual competitive pressure of all neighbors in a single competitive pressure vector, we were able to predict the direction of crown asymmetry of the focal tree with an accuracy of 96° on the full circle (360°).The competitive pressure model was equally applicable to beech, ash and hornbeam trees and may generate valuable insight into competitive interactions among tree crowns in mixed stands, provided that sufficiently precise data on the shape and position of the tree crowns is available. Multiple-aspect laser-scanning proved to be an accurate and practicable approach for analyzing the complex 3-dimensional shape of the tree crowns, needed to quantify the plasticity of growth processes in the canopy. We conclude that the laser-based analysis of crown plasticity offers the opportunity to achieve a better understanding of the dynamics of canopy space exploration and also may produce valuable advice for the silvicultural management of mixed stands.     

Modeling radiation transfer within the canopy of a Chinese fir plantation

Calculation of radiation transfer within the canopy of a plantation on sloped sites is described. The canopy was assumed to consist of evenly cone-shaped crowns. Within-tree shading by object tree and between-tree shading by tree crowns around the object tree were differentiated. The two-dimensional needle area density (NAD) was introduced in the calculation of radiation transfer and interception. The model was validated with measurements of both daily photosynthetic photo flux density (PPFD) and daily photosynthetic radiation at tens of points within the crown of a 17-year-old Chinese fir (Cunninghamia lanceolata) plantation growing on a sloped site. Simulation showed that modeled data closely matched measurements. Simulation errors likely result from the uneven size of tree crowns, irregular crown shape, grouping of needles to shoots and twigs, etc. Scattering radiation amounted to below 10%, in most cases, of daily radiation, and its uncertainty was relatively small.     

Vegetation and weather explain variation in crown damage within a large mixed-severity wildfire

The 2002 Biscuit Fire burned through more than 200,000 ha of mixed-conifer/evergreen hardwood forests in southwestern Oregon and northwestern California. The size of the fire and the diversity of conditions through which it burned provided an opportunity to analyze relationships between crown damage and vegetation type, recent fire history, geology, topography, and regional weather conditions on the day of burning. We measured pre- and post-fire vegetation cover and crown damage on 761 digital aerial photo-plots (6.25 ha) within the unmanaged portion of the burn and used random forest and regression tree models to relate patterns of damage to a suite of 20 predictor variables. Ninety-eight percent of plots experienced some level of crown damage, but only 10% experienced complete crown damage. The median level of total crown damage was 74%; median damage to conifer crowns was 52%. The most important predictors of total crown damage were the percentage of pre-fire shrub-stratum vegetation cover and average daily temperature. The most important predictors of conifer damage were average daily temperature and “burn period,” an index of fire weather and fire suppression effort. The median level of damage was 32% within large conifer cover and 62% within small conifer cover. Open tree canopies with high levels of shrub-stratum cover were associated with the highest levels of tree crown damage, while closed canopy forests with high levels of large conifer cover were associated with the lowest levels of tree crown damage. Patterns of damage were similar within the area that burned previously in the 1987 Silver Fire and edaphically similar areas without a recent history of fire. Low-productivity sites on ultramafic soils had 92% median crown damage compared to 59% on non-ultramafic sites; the proportion of conifer cover damaged was also higher on ultramafic sites. We conclude that weather and vegetation conditions — not topography — were the primary determinants of Biscuit Fire crown damage.     

Tree allometry, leaf size and adult tree size in old-growth forests of western Oregon

Relationships between tree height and crown dimensions and trunk diameter were determined for shade-tolerant species of old-growth forests of western Oregon. The study included both understory and overstory species, deciduous and evergreen angiosperms and evergreen conifers. A comparison of adult understory species with sapling overstory species of similar height showed greater crown width and trunk diameter in the former, whether the comparison is made among conifers or deciduous trees. Conifer saplings had wider crowns than deciduous saplings, but the crown widths of the two groups converged with increase in tree height. Conifer saplings had thicker trunks than deciduous saplings of similar crown width, possibly because of selection for resistance to stem bending under snow loads. The results suggest that understory species have morphologies that increase light interception and persistence in the understory, whereas overstory species allocate their biomass for efficient height growth, thereby attaining the high-light environment of the canopy. The greater crown widths and the additional strength requirements imposed by snow loads on conifer saplings result in less height growth per biomass increment in conifer saplings than in deciduous saplings. However, the convergence in crown width of the two groups at heights greater than 20 m, and the proportionately smaller effect of snow loads on large trees, may result in older conifers equalling or surpassing deciduous trees in biomass allocation to height growth.     

Forest shadows: How much shelter in a shelterwood?

A simulation model is presented that will permit silviculturists to estimate the shade cast on any site at any time by a randomly distributed conifer stand of any stocking level. Computational requirements are modest. Simple situations may be solved in an hour or so on a programmable hand calculator; most complex situations can be handled on a desk microcomputer.Trees are modelled as a crown consisting of a cone on top of a cylinder, and the crowns may vary in density. The bole is modelled as a solid cylinder. Relative dimensions may be varied to simulate almost all regular conifer tree forms. The projected shadow area of trees is integrated for any number od independently distributed trees to obtain the relative shadow area on the ground surface. Alternatively, the model permits the determination of the number of trees of any given dimensions per unit are needed to provide a given degree of shade.     

The effects of Douglas-fir on tree-specific arthropod communities in mixed species stands with European beech and Norway spruce

The ecological effects of planting exotic Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] in Central Europe are still poorly understood. The aim of this study was to answer the question of whether Douglas-fir affects tree specific arthropod communities in different mature forest types (Douglas-fir, spruce and beech dominated) in Southern Germany. Therefore, arthropod communities of stem and tree crown strata of Douglas-fir and spruce (Picea abies L.) were sampled in the years 1999–2001 using arboreal photo-eclectors and flight interception traps. Statistical analysis was conducted for all species and focused on conifer specialists at three levels: (1) species diversity, (2) guild structure and (3) community structure. Within the stem stratum, species diversity was significantly higher on spruce than on Douglas-fir independent of year and stand composition. This could not be explained by a single feeding guild, rather by species changing strata during the vegetation period. In contrast, species diversity in tree crowns was approximately the same for both conifer species. However, communities in Douglas-fir crowns were conspicuously different from those in spruce crowns, especially in the Douglas-fir dominated stand type. While zoophagous insects exhibited higher activity on Douglas-fir in 2000, xylophagous beetles were more abundant on spruce in 2001. In European beech stands with widely spaced Douglas-fir trees, the site specific and broad-leaved tree related fauna might be maintained. In addition, Douglas-fir with its resource of Adelges cooleyi and crowns that overtop the broad-leaved tree canopy, offer additional resources for several aphidophagous and thermophile species.     

Liana abundance in a Puerto Rican forest

Liana (woody vine) abundance varies among tropical forests and is often high in disturbed forests. In two areas of subtropical wet forest in Puerto Rico, El Verde and Bisley, we recorded the density of liana stems ≥1 cm dbh, and the percent of tree crowns (trees ≥10 cm dbh) that lianas infested. Both study areas have been disturbed by hurricanes several times in the past century; however, sample plots in each area were divided between plots that were less disturbed and those that were more disturbed, by both hurricanes and humans. The mean density and basal area of liana stems at El Verde were significantly higher in the less disturbed plots than in the more disturbed plots. The percent tree crown infested by lianas was higher on certain tree species and on larger trees, both of which characterized the less disturbed forest. Results at Bisley were similar to those at El Verde. Liana density and tree crown infestation in these Puerto Rican forests were low compared with most other tropical forests, contrasting especially with high values in other disturbed forests. Liana abundance varies among forests for complex reasons, including differences in disturbance, biogeography, seasonality, and tree host features.     

The effects of belowground resources on aboveground allometric growth in Bornean tree species

Tree height and crown allometries reflect adaptations for resource acquisition and structural stability, as well as plastic responses to a heterogeneous environment. While both light and soil resources limit growth and influence competitive responses in tropical forests, the effects of belowground resources on allometries are less understood, especially within the understory. To characterize outcomes of tree competition along an edaphic resource gradient, we quantified variation in height and crown allometries of six Bornean tree species from contrasting regeneration niches (light-demanding vs. shade-tolerant) on two soil habitats (clay-fine loam and sandy loam) within a 52-ha forest dynamics plot. Using empirically-fit allometric parameters and diameter growth rates from plot census data, we modeled tree height and crown area growth over the projected life span of each species. Based on resource competition theory, we hypothesized that tree species specializing on and populations of generalist species growing on the relatively moister, more fertile clay-fine loam soil habitat would have faster height and crown growth rates, compared to those on the sandy loam habitat, regardless of regeneration niche. Among soil specialists and within generalists of both genera, trees growing on clay-fine loam had taller stems and larger crowns at a given age and faster height and crown area growth rates at most sizes than trees on sandy loam. Differences in height and crown growth were driven by the faster diameter growth rates of trees on clay-fine loam, not by differences in height- and crown-diameter allometries, as trees on sandy loam were significantly taller at a given diameter, and differences in crown allometry were not consistent across soil habitats. Characterizing the height and crown growth responses of trees along resource gradients provides insight into the mechanisms that maintain diversity in tropical forests. Our results point to the importance of adaptive and plastic responses to both above and belowground resource availability in determining the allometric growth of trees and suggest that this diversity of responses may contribute tree species coexistence through competition-based trade-off mechanisms and variation in growth among individuals. Additionally, as the importance estimating natural carbon sequestration increases with the escalating effects of anthropogenic climate change, differences in tree growth and architecture across soil habitats also have implications for the approximation of forest carbon storage on heterogeneous tropical soils.     

Hollows in jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) trees: I. Hollow sizes, tree attributes and ages

One hundred and fifty-four jarrah (Eucalyptus marginata) and 85 marri (Corymbia calophylla) trees were measured and assessed, and the numbers and sizes of hollows in these trees were determined by destructive sampling; 665 hollows were located and measured. The relationship between tree diameter and tree age was determined from counts of annual growth rings on 162 of these trees. Large trees and trees with moderately senescent crowns individually bear the most hollows. Although the number of hollows found in individual trees increased with tree diameter, the distribution of tree diameters in the forest is skewed and the large number of small trees with diameters between 40 and 80 cm contribute approximately 50% of all hollows in the jarrah forest. The distributions of entry size, and of hollow depth, are highly skewed, with small hollows occurring more frequently than large hollows. Although jarrah trees bear more hollows than marri trees and the distribution of entry sizes is similar for both tree species, the hollows in jarrah are significantly smaller than the hollows in marri. Most hollows are cylindrical in shape, vertically oriented and occur in dead wood in the tree crown. Relatively few hollows (14%) occur in the tree bole or at crown break. Counts of hollows made from ground level are inaccurate as estimates of the actual number of hollows in trees.     

Applied 3D texture features in ALS-based forest inventory

Airborne laser scanning (ALS) data are not usually considered to be very informative with respect to tree species, and this information is often obtained by combining such data with spectral image material. The aim was to test the ability of height, density, intensity and applied 2D and 3D texture variables derived solely from a very high-density ALS point cloud to describe the crown shape and structure characteristics required for tree species discrimination. Linear discriminant analysis was used to find optimal combinations of variables within the predictor groups, and classifications based on variables from different groups were compared. The third power of the tree diameter was used as a stem volume approximate, and rather than examining species alone, the classification was evaluated with respect to the volume approximates assigned to the predicted species. The sensitivity of pulse density to the methodology presented here was determined by simulating thinned data sets by reducing the initial pulse density. The reliability of the estimates was analysed both with functions generated using the original data and with new functions for each thinning level. Alpha shape metrics developed for describing tree crowns constructed from the 3D point clouds proved capable of discriminating between all three species groups evaluated, and several height distribution and textural variables were found to discriminate between the coniferous tree species. The results demonstrate the importance of species interpretation in forest inventories based on allometric modelling, but then indicate that species-specific estimation could be carried out using ALS-derived variables alone.     

Estimation of aboveground biomass in mangrove forests using high-resolution satellite data

Mangroves play important roles in providing a range of ecosystem services, mitigation of strong waves, protection of coastlines against erosion, maintenance of water quality, and carbon sink in the context of global warming. For trees in mangrove forests in southern Ranong Province, Thailand, we investigated the allometric relationship between crown area derived from high-resolution satellite data and stem diameter and used the resulting model to estimate aboveground biomass. We used QuickBird panchromatic and multispectral data acquired for the study area on 15 October 2006 as the high-resolution satellite data. Individual tree crowns were extracted from the satellite image of panchromatic data by using the watershed method, and the species were identified by using the maximum-likelihood method for the multispectral data. Overall classification accuracy for species identification was 88.5 %. The biomass derived from our field survey was plotted against aboveground biomass in the sample plots, estimated from the QuickBird data. The regression line through the origin between the satellite-estimated biomass and biomass based on the field data had a slope of 1.26 (R ² = 0.65). Stand aboveground biomass estimated from the high-resolution satellite data was underestimated because of a lack of data on the biomass of suppressed trees and inappropriate segmentation of crowns of large trees into two or more trees.     

Millet production under pruned tree crowns in a parkland system in Burkina Faso

As a tree management tool, three treatments of crown pruning (total-pruning, half-pruning and no-pruning) were applied to Vitellaria paradoxa (karité) and Parkia biglobosa (néré) in agroforestry parkland systems in Burkina Faso. The area under each tree was divided into four concentric tree influence zones (Zones A: up to 2 m from the tree trunk, B: up to half of the radius of the tree crown, C: up to the edge of the tree crown and D: up to 2 m away from the edge of the tree crown). Millet production under these zones and outside was assessed during two cropping seasons over the study period of three years and the results showed that tree crown pruning had significant effect on millet production and the highest millet grain yield and total dry matter were produced under total-pruned trees (507 ± 49 and 2033 ± 236 kg ha⁻¹ year⁻¹, respectively). Light transmission, transpiration and soil nutrient status under the trees were also analysed in relation to millet production. The results of the analysis showed that total-pruned trees gave the highest millet production due to the reduction by crown pruning of the effects of large tree crowns on PAR transmission below crowns and rates of transpiration by trees. Soil was more fertile closer to the tree trunks than outside tree crowns. This may also be one of the reasons why millet overall performed better under Zone B than outside tree crowns. The higher production of millet under Zone B than under Zone A, the zone closer to the tree trunk, may be due to lower light intensity and more intense competition for water between trees and crops under Zone A. It was concluded that at least in the short term millet production could be improved by crown pruning of both karité and néré, but long term effects may depend on the ability of the trees to maintain the amelioration of soil fertility and on how quickly the trees recover from pruning. This revised version was published online in June 2006 with corrections to the Cover Date.     

基于大比例尺航片的针叶树种冠幅的提取

基于凉水国家级自然保护区2009年拍摄的1：2000航空像片和同期森林资源二类调查的固定样地数据,采用子像元分类方法分别提取出红松、落叶松和云冷杉的专题影像图。在此基础上,将栅格专题影像图转换为矢量图形,采用目视解译的方法提取上层针叶林的树冠信息。通过将针叶树冠形似为圆形提取出各树种的冠幅,用固定样地实测数据进行对比分析和精度评价,并建立航片上提取冠幅与实测冠幅之间的一元线性回归模型。结果表明：红松、落叶松和云冷杉冠幅的提取精度分别达到83．50％、84．35％和82．26％,其预测精度分别达到83．60％、81．46％和83．57％。     

Determination of the spatial distribution of trees from digital aerial photographs

This study examined the possibilities of using computerized digital aerial photograph interpretation in determining the spatial distribution of trees. The material of the study included eight mapped stands in the municipality of Hyytiälä (61°50′N and 24°18′E), in southern Finland. The aerial photographs used were taken in June 1995 at a scale of 1:5000. Two approaches for determining the spatial pattern of trees were used. Firstly, in the point-process based approach used in this study, the individual trees in the digital aerial photograph were segmented by a robust segmentation method, based on recognition of the pattern of tree crowns with sub-pixel accuracy. Secondly, the crown coverage was determined by region growing segmentation combined with active surface representation. The significance of the differences in the means of image coverage pattern indices between the various spatial distribution categories was tested with one-way variance analysis. Because the process misclassified clustered spatial patterns as regular patterns, and regular patterns as random patterns, the usability of digital aerial photographs seems to be limited for the point-process based determination of the spatial pattern of trees if the scale is 1:5000 or less. When image coverage pattern indices were applied, the differences in the means of the spatial distribution categories proved not to be clearly statistically significant due to the great variation within classes. However, interpretation of crown coverage could have applications in practical forestry due to the low resolution requirements for the images used.     

Hollows in jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) trees: II. Selecting trees to retain for hollow dependent fauna

From a sample of 665 hollows found in 154 jarrah (Eucalyptus marginata) and 85 marri (Corymbia calophylla), we identified 204 hollows in 84 trees that were potentially suited to one or more of 10 species of hollow using birds and mammals. Occurrence of these hollows increased with tree age, tree size, and species (marri bore more usable hollows than jarrah) and increased amounts of dead wood in tree crowns. Hollow occurrence was most likely in trees with moderately senescent crowns with damage to intermediate sized branches, and the largest hollows were more likely to occur in more highly senescent crowns. Evidence of termite invasion at the tree butt was not related to occurrence of hollows. For all but one of the birds and mammals we considered, dead trees were no more likely than live trees to contain hollows.

Our study indicates that for the purpose of forest management planning, 130 years can be taken as the typical minimum age for the formation of usable hollows in jarrah and marri. The current minimum prescribed diameter for “habitat trees” (trees retained in logged areas to supplement existing hollows), which corresponds to a mean age of 171 years, is thus a realistic minimum size for these retained trees. We recommend raising the prescribed range of crown senescence for retained habitat trees to increase the probability of providing large hollows suited to large species such as red-tailed black cockatoo, and common brushtail possum, and maternal hollows used by smaller species. Retaining the largest trees with appropriate crown attributes will substantially increase the probability that these trees will bear usable hollows.     

The contribution of structural indices to the modelling of Sitka spruce (Picea sitchensis) and birch (Betula spp.) crowns

Crown dimensions are important for the quantification of tree interactions in some growth models. This study investigates the potential for structural indices and other spatial measures to improve the prediction of crown radius and crown length for birch (Betula spp.) and Sitka spruce (Picea sitchensis (Bong.) Carr.) in forests in Wales. Crown dimensions were measured for 125 birch and 154 spruce in six fully stem-mapped research plots. These data were used to test the performance of a crown radius model and a crown length model which estimated crown dimensions on the basis of allometric relationships with stem dimensions. Spatial data from the six plots were used to calculate the structural indices mean directional index, diameter correlation index, species mingling, dbh and height dominance, and dbh differentiation, as well as the Hegyi competition index, and basal area of neighbours and larger neighbours, for each crown measurement sample tree, using various numbers of nearest neighbours. Two non-spatial indices, BAL and BALMOD, were also calculated for all sample trees for comparison. These spatial and non-spatial variables were then incorporated into modified crown dimension models. Model performances, in terms of efficiency and relative bias, were compared to determine whether the inclusion of spatial or non-spatial variables resulted in any improvements over models using tree dimensions alone. Crown length and radius were found to be correlated with most of the spatial measures studied. Models incorporating spatial variables gave improvements in performance over allometric models for every data set, and performed more consistently than models containing non-spatial variables. The greatest improvements were achieved for suppressed birch in unthinned forests which had irregularly shaped and strongly displaced crowns. The spatial variable contributing to the most efficient model for each data set varied widely. This points to the complexity of tree spatial interactions and indicates that there is a great deal of scope for investigating other structural indices and crown dimension model forms.     

Preformation in vegetative buds of pistachio (Pistacia vera): relationship to shoot morphology, crown structure and rootstock vigor

Effects of rootstock, shoot carbohydrate status, crop load and crown position on the number of preformed leaf primordia in the dormant terminal and lateral buds of mature and immature 'Kerman' pistachio (Pistacia vera L.) trees were investigated to determine if rootstock vigor is associated with greater shoot preformation. There was no significant variation in preformation related to the factors studied, suggesting strong genetic control of preformation in 'Kerman' pistachio. The growth differences observed among trees on different rootstocks were associated with greater stimulation of neoformed growth in trees on the more vigorous rootstocks. However, most annual extension growth in mature tree crowns was preformed, contrasting with the relatively high rate of neoformation found in young tree crowns. Large amounts of neoformed growth in young trees may allow the trees to become established quickly and secure resources, whereas predominantly preformed growth in mature trees may allow for continued crown expansion without outgrowing available resources. We hypothesized that the stimulation of neoformed growth by the more vigorous rootstocks is associated with greater resource uptake or transport, or both. Understanding the source of variation in shoot extension growth on different rootstocks has important implications for orchard management practices.     

Multiple attribute decision making for individual tree detection using high-resolution laser scanning

A canopy height model (CHM) is a standard LiDAR-derived product for deriving relevant forest inventory information, including individual tree positions, crown boundaries and plant density. Several image-processing techniques for individual tree detection from LiDAR data have been extensively described in literature. Such methods show significant performance variability depending on the vegetation characteristics of the monitored forest. Moreover, over regions of high vegetation density, existing algorithms for individual tree detection do not perform well for overlapping crowns and multi-layered forests. This study presents a new time and cost-efficient procedure to automatically detect the best combination of the morphological analysis for reproducing the monitored forest by estimating tree positions, crown boundaries and plant density from LiDAR data. The method needs an initial calibration phase based on multi attribute decision making-simple additive weighting (MADM-SAW). The model is tested over three different vegetation patterns: two riparian ecosystems and a small watershed with sparse vegetation. The proposed approach allows exploring the dependences between CHM filtering and segmentation procedures and vegetation patterns. The MADM architecture is able to self calibrate, automatically finding the most accurate de-noising and segmentation processes over any forest type. The results show that the model performances are strongly related to the vegetation characteristics. Good results are achieved over areas with a ratio between the average plant spacing and the average crown diameter (TCI) greater than 0.59, and plant spacing larger than the remote sensing data spatial resolution. The proposed algorithm is thus shown a cost effective tool for forest monitoring using LiDAR data that is able to detect canopy parameters in complex broadleaves forests with high vegetation density and overlapping crowns and with consequent significant reduction of the field surveys, limiting them over only the calibration site.     

Carbohydrate transfer through root grafts to support shaded trees

We investigated whether root grafts between lodgepole pine (Pinus contorta var. latifolia Dougl. ex. Loud.) trees can transfer sufficient carbohydrate reserves from a source tree to a grafted sink tree to affect the vigor of trees growing in a light-limited environment. Eleven plots were established in early spring and two grafted tree pairs and two independent non-grafted trees were selected at each plot. One tree in a grafted pair and one non-grafted tree were shaded at each plot, whereas the remaining trees were non-shaded during the experimental period. Shaded trees had significantly lower carbohydrate reserves and smaller crowns than non-shaded trees following one growing season. Grafted shaded trees had significantly higher root total nonstructural carbohydrate concentrations than non-grafted shaded trees, indicating that root grafts partially offset the effects of shading. Also, large root grafts transferred proportionately more carbohydrates to the shaded trees than small root grafts. Carbohydrates transferred through root grafts could allow grafted trees to persist under conditions where non-grafted trees would be removed by competition.     

Coordination of crown structure, leaf plasticity and carbon gain within the crowns of three winter-deciduous mature trees

We examined the vertical profiles of leaf characteristics within the crowns of two late-successional (Fagus crenata Blume and Fagus japonica Maxim.) and one early-successional tree species (Betula grossa Sieb. et Zucc.) in a Japanese forest. We also assessed the contributions of the leaves in each crown layer to whole-crown instantaneous carbon gain at midday. Carbon gain was estimated from the relationship between electron transport and photosynthetic rates. We hypothesized that more irradiance can penetrate into the middle of the crown if the upper crown layers have steep leaf inclination angles. We found that such a crown has a high whole-crown carbon gain, even if leaf traits do not change greatly with decreasing crown height. Leaf area indices (LAIs) of the two Fagus trees (5.26-5.52) were higher than the LAI of the B. grossa tree (4.50) and the leaves of the F. crenata tree were more concentrated in the top crown layers than were leaves of the other trees. Whole-crown carbon gain per unit ground area (micromol m(-2) ground s(-1)) at midday on fine days in summer was 16.3 for F. crenata, 11.0 for F. japonica, and 20.4 for B. grossa. In all study trees, leaf dry mass (LMA) and leaf nitrogen content (N) per unit area decreased with decreasing height in the crown, but leaf N per unit mass increased. Variations (plasticity) between the uppermost and lowermost crown layers in LMA, leaf N, the ratio of chlorophyll to N and the ratio of chlorophyll a to b were smaller for F. japonica and B. grossa than for F. crenata. The light extinction coefficients in the crowns were lower for the F. japonica and B. grossa trees than for the F. crenata tree. The leaf carbon isotope ratio (delta(13)C) was higher for F. japonica and B. grossa than for F. crenata, especially in the mid-crown. These results suggest that, in crowns with low leaf plasticity but steep leaf inclination angles, such as those of F. japonica and B. grossa trees, irradiance can penetrate into the middle of the crowns, thereby enhancing whole-crown carbon gain.