Leaf mass per area relationships across light gradients in hybrid spruce crowns

This study examined the distribution and variation of mass to projected area ratio of foliage (LMA, g/m²) within hybrid spruce (Picea engelmannii Parry×Picea glauca (Moench) Voss×Picea sitchensis (Bong.) Carr) crowns across three stages of stand development (20, 60, and 140 years). Variability of LMA was assessed at different heights and branch positions. LMA decreased with distance from top of tree (p<0.0001) at rates that varied among stand development stages (p<0.0001). A multi-level mixed effect analysis indicated that distance from the tip of the first-order branch (p=0.0002) had a significant influence on LMA. In general, LMAs decreased towards the base of the tree and increased towards tree apex.LMA differed significantly across stand development stages. Older trees (140 years) showed the highest LMA, while younger trees (20 years) showed the lowest LMA values. LMA also increased with foliage age, suggesting a developmental change in leaf area and mass with increasing foliage age. Multiple linear regression (MLR) relationships were developed to predict LMA at various positions within tree crowns. The precision of the models was slightly greater when branch position (from the apex of the tree) was described relative to tree height, rather than relative to crown length. The MLR function resulted in precise representations of LMA within tree crowns.     

Variation of specific leaf area and upscaling to leaf area index in mature Scots pine

Reliable and objective estimations of specific leaf area (SLA) and leaf area index (LAI) are essential for accurate estimates of the canopy carbon gain of trees. The variation in SLA with needle age and position in the crown was investigated for a 73-year-old Scots pine (Pinus sylvestris L.) stand in the Belgian Campine region. Allometric equations describing the projected needle area of the entire crown were developed, and used to estimate stand needle area. SLA (cm² g⁻¹) as significantly influenced by the position in the crown and by needle age (current-year versus 1-year-old needles). SLA increased significantly from the top to the bottom of the crown, and was significantly higher near the interior of the crown as compared to the crown edge. SLA of current-year needles was significantly higher than that of 1-year-old needles. Allometric relationships of projected needle area with different tree characteristics showed that stem diameter at breast height (DBH), tree height and crown depth were reliable predictors of projected needle area at the tree level. The allometric relationships between DBH and projected needle area at the tree level were used to predict stand-level needle area and estimate LAI. The LAI was 1.06 (m² m⁻²) for current-year needles and 0.47 for 1-year-old needles, yielding a total stand LAI of 1.53.     

Crown profile of foliage area characterized with the Weibull distribution in a hinoki (Chamaecyparis obtusa) stand

Summary Thirten sample trees of various sizes in a 29-year-old hinoki [Chamaecyparis obtusa (Sieb, et Zucc.) Endl.] plantation were felled and subjected to the stratified clip technique. Crown profile of foliage area fitted well with the Weibull distribution. The crown profile tended to be more skewed toward the top of crowns in smaller trees than in larger trees. This tendency was reflected in the value of the shape parameter of the Weibull distribution. The shape parameter ranged from 1.73 to 3.23 and gradually increased up to an asymptotic value with an increase of stem diameter at breast height. The scale parameter of the distribution ranged from 1.0 to 4.2 and tended to increase in proportion to stem diameter at breast height. Foliage area of a tree correlated well with stem diameter at breast height through an ordinary allometric equation. Tree height could be approximated fairly well by a generalized allometric equation as a function of stem diameter at breast height. On the basis of the census of stem diameter at breast height, canopy profile could be constructed synthesizing crown profiles of foliage area for individual trees in the stand. Leaf area index was estimated to be 6.6 ha ha^–1.     

Variation in crown light utilization characteristics among tropical canopy trees

BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.     

Patterns of Trunk Diameter, Tree Height and Crown Depth in Crowded Abies Stands

The size structure of trees in crowded, even-aged Abies (fir)stands of ‘Shimagare’ or ‘wave-regenerated’sub-alpine forests is analyzed. Tree-height distributions showconsistently smaller variation and less positive skewness thanthe distributions of trunk diameter and crown depth (tree heightminus height of the lowest branch). This difference is associatedwith changes in the relationships between trunk diameter, treeheight and crown depth as stands age. These, in turn, resultfrom self-pruning of the lower foliage crown due to competitionfor light in crowded stands. Abies, diameter-height curve, competition, size distribution, stand development, tree geometry, wave-regeneration     

Stomatal activity within the crowns of tall deciduous trees under forest conditions

The variation in stomatal activity within the crowns ofAcer campestre, Carpinus betulus andQuercus cerris was measured by vapour exchange porometer on several summer days in an oak-hornbeam forest, in SW Slovakia, Czechoslovakia. Variation resulted from crown position in the forest stand and from leaf position within the canopy. The highest stomatal conductance was in sunlit sun leaves in the upper part of the canopy. Stomatal conductance decreased with increasing depth in the canopy. The steepest decrease was in the upper canopy, in the intermediate zone between fully sunlit and fully shaded leaves, and was caused by the decline in leaf irradiance and in stomatal density. In codominant trees, the conductance in shade leaves at the base of the crown was significantly lower than in the sun leaves at the top of the crown. In a dominant tree,Q. cerris, the differences in stomatal conductance were small and most frequently insignificant. Variation in incident light also determined the diurnal variation of stomatal conductance with respect to crown aspect. Differences between sun leaves on the east and west facing aspects of the overstory crown ofQ. cerris were demonstrated for several days.     

Changes in foliage distribution with relative irradiance and tree size: Differences between the saplings ofAcer platanoides andQuercus robur

Dependencies of foliage arrangement and structure on relative irradiance and total height (TH) were studied in saplings ofAcer platanoides andQuercus robur. The distribution of relative foliar area and dry weight (leaf area and weight in a crown layer per total tree leaf area and weight, respectively) were examined with respect to relative height (RH, height in the crown per TH) and characterized by the Weibull function. The distributions of relative area and weight were nearly identical, and the differences between them were attributable to a systematic decline in leaf dry weight per area with increasing crown depth. Foliage distribution was similarly altered by tree size in both species; RH at foliage maximum was lower and relative canopy size (RCS, length of live crown per TH) greater in taller trees. However, the distribution was more uniform inA. platanoides than inQ. robur. Apart from the size effects, relative irradiance also influenced canopy structure; RCS increased inQ. platanoides and decreased inQ. robur with increasing irradiance. As crown architecture was modified by irradiance, foliage distribution was shifted upwards with decreasing irradiance inA. platanoides, but it was independent of irradiance inQ. robur. Higher foliage maximum at lower irradiance in more shade-tolerantA. platanoides is likely to contribute towards more efficient foliar display for light interception and increase the competitive ability of this species in light-limited environments. Consequently, these differences in crown architecture and foliage distribution may partly explain the superior behavior ofA. platanoides in understory.     

Competition-dependent modelling of foliage biomass in forest stands

Currently, foliage biomass is estimated based on stem diameter or basal area. However, it is questionable whether the relations between foliage and stem observed from plantations of a single tree species can be applied to stands of different structure or species composition. In this paper, a procedure is presented to simulate foliage and branch biomass of tree crowns relative to crown size and light competition. Crowns are divided into layers and segments and each segment is divided into a foliated and an unfoliated fraction. Depending on the competitive status of the segment, leaf area density, specific leaf area and foliated branch fraction are determined. Based on this information, foliage biomass is calculated. The procedure requires a crown shape function and a measure to characterise competition for light and space of each individual segment within the canopy. Simple solutions are suggested for both requirements to enable an application with data that can be measured non-destructively in the field; these were stem position, tree height, crown base height, crown radii and some general crown shape information. The model was parameterised from single trees of Norway spruce and European beech and partly evaluated with independent data close to the investigation plot. Evaluations showed that the model can attribute the ecology of the different crown forms. Modelled foliage distribution for beech and spruce as well as total needle biomass of spruce agreed well with measurements but foliage biomass of beech was underestimated. The results are discussed in the context of a general model application in structured forests.     

Comparative height crown allometry and mechanical design in 22 tree species of Kuala Belalong rainforest, Brunei, Borneo

In rainforests, trunk size, strength, crown position, and geometry of a tree affect light interception and the likelihood of mechanical failure. Allometric relationships of tree diameter, wood density, and crown architecture vs. height are described for a diverse range of rainforest trees in Brunei, northern Borneo. The understory species follow a geometric model in their diameter-height relationship (slope, β = 1.08), while the stress-elasticity models prevail (β = 1.27-1.61) for the midcanopy and canopy/emergent species. These relationships changed with ontogeny, especially for the understory species. Within species, the tree stability safety factor (SSF) and relative crown width decreased exponentially with increasing tree height. These trends failed to emerge in across-species comparisons and were reversed at a common (low) height. Across species, the relative crown depth decreased with maximum potential height and was indistinguishable at a common (low) height. Crown architectural traits influence SSF more than structural property of wood density. These findings emphasize the importance of applying a common reference size in comparative studies and suggest that forest trees (especially the understory group) may adapt to low light by having deeper rather than wider crowns due to an efficient distribution and geometry of their foliage.     

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.     

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.     

Angle distribution of foliage in individual Chamaecyparis obtusa canopies and effect of angle on diffuse light penetration

 The vertical distribution of foliage angle and area of three Chamaecyparis obtusa trees was determined by the triangle method, which calculates foliage geometry using measured coordinates of the leaf ”corners”, in a 43-year-old plantation in central Japan. Vertical distribution patterns of leaf area were different depending on tree size, but the boundary heights, which divide the canopy into sunlit and shaded parts, were similar in the three sample trees. The value of the average foliage angle [I(Z)] at a given depth (Z) from the tip of the stem decreased continually from the upper to lower layers within the canopy. The vertical patterns of changes in I(Z) were different among the three trees, but could be expressed by the following allometric equation as a function of depth.

Causes of vertical stratification in the density of Cameraria hamadryadella

1. The density of Cameraria hamadryadella, a leaf-mining moth, is vertically stratified within the crown of oak trees. It occurs at higher densities on foliage in the lower crown. 2. Oviposition preference tests indicate that females display no preference to oviposit on foliage from the lower tree crown over foliage from the upper tree crown. 3. Experiments in which potted trees were placed at various heights indicate that foliage nearest ground level receives more oviposition, and that the higher rates of oviposition on foliage near the ground is not caused by differences in quality between foliage from low or high in the tree crown. 4. Host-plant- and natural-enemy-mediated juvenile mortality and the abundance of parasitoids did not differ between the upper and lower crown. 5. Vertical differences in the timing of leaf production within the tree crown are unlikely to account for the observed gradient in the abundance of C. hamadryadella. Furthermore, the mean date of leaf fall does not differ among heights within the tree crown. 6. It is argued that microclimatic gradients and interspecific competition are also unlikely to account for the observed gradients in the density of C. hamadryadella within the tree crown. 7. Because of the absence of effects of other potentially causal factors, the most likely explanation for the gradient in density of C. hamadryadella is a lack of movement by females into the upper tree crown from overwintering, emergence, or resting sites located in the lower tree crown, combined with egg depletion by females as they move from the lower to the upper tree crown. 8. It is suggested that the lack of movement and egg depletion hypothesis should serve as the null hypothesis in studies of vertical stratification of tree-feeding insects. In the absence of evidence of an effect of foliage quality, natural enemies, plant phenology, interspecific competition, or microclimate, the movement and egg depletion hypothesis is the most reasonable.     

北京地区侧柏人工林密度效应

密度是影响森林尤其是人工林生长的重要因素,林冠层是森林生态系统与其他系统进行能量和物质交换的重要场所,树木及树冠生长对林分密度的响应关系可以看作是生物对环境变化产生的适应性现象。林分密度效应是生态学和森林培育学的重要研究内容之一。以23块8种不同密度梯度的北京山区侧柏人工幼龄林林分为研究对象分析其树木生长及树冠生长对密度的响应关系,其中树冠指标使用了参照了美国林务局(USDA)的树冠调查指标。研究结果表明:(1)林分平均胸径、平均树高和平均冠幅生长均随密度增大而减小,林分密度大于3000株/hm2时各指标减小的趋势变缓,使用异速生长模型可以很好地拟合这种变化关系;(2)随密度增加,树冠水平方向和垂直方向生长均到显著地抑制作用,树冠外形表现出由饱满冠型向狭长冠型变化的适应性现象;(3)使用树冠二维、三维指标与密度进行相关性分析可知树冠长度、树冠率等指标与林分密度呈负相关关系,树冠圆满度及树冠生产效率与密度表现出极显著正相关关系;(4)采用枝解析的方法研究了树枝长度、材积的平均生长量、连年生长量与密度的关系,结果表明幼龄期各生长量差异不大;(5)在建立冠幅模型时考虑了自变量间的多重共线性问题,所建的胸径单自变量二次方模型能够很好地预测侧柏人工幼龄林冠幅生长过程,模型相关系数R2为0.961。     

Crown structure and leaf area of the understorey species Prunus serotina

A detailed biometrical study of the exotic understorey invader Prunus serotina (Ehrh.) was carried out in a mixed coniferous forest stand in northern Belgium. Based on detailed destructive measurements of eight selected model trees, allometric relations of tree height, crown projected area, woody and leaf dry mass and leaf area on tree diameter at breast height (DBH) were derived. The scaling-up procedure from the tree to the stand level was done using the frequency distribution of DBH obtained at the selected experimental plot. The vertical and radial distributions of the tree foliage were estimated by the “cloud” technique. The vertical profile of leaf area showed a bimodal distribution pattern with maxima at heights of 4 and 6 m above the ground. The leaf area index (LAI) of the understorey Prunus serotina as estimated by the described up-scaling procedure (5.1) was significantly higher than the LAI (2.6) as measured by a plant canopy analyser and was also significantly higher than the LAI of the overstorey species Scots pine (1.5–3.0). The LAI of a neighbouring Rhododendron understorey reached only 1.25. This study emphasises the importance of an exotic understorey species in the total leaf area of mixed coniferous forests which might have important implications for the energy and mass exchanges of the entire forest.     

Crown development in Norway spruce [<Emphasis Type="Italic"> Picea abies </Emphasis>(L.) Karst.]

This study tests whether crown and stem development in Norway spruce could be described using a modified profile theory. 29 trees from three age-groups (25, 67, 86) with different treatments (unthinned, normally and intensively thinned) were destructively sampled. Crown ratio and crown length varied between age groups and treatments. Crown width was positively correlated with crown length, but branch length along the crown depended on tree age and growing space. Foliage mass density peaked at a relative crown height of 50–70% in middle-aged and mature stands, while young crowns were densest and widest at the base. Foliage mass was predictable from branch and stem cross-sectional area, provided the distance from the top was included. The ratio of foliage mass to branch cross-sectional area increased for 2–4 m down from the tip of the crown, then started to decrease. The relationship between cumulative foliage mass and stem cross-sectional area was non-linear along the stem in the upper crown, but the ratio of cumulative branch to stem cross-sectional area was linear. Trees in the mature and unthinned stands had more cross-sectional area in branches relative to stems than in the young and thinned stands. We conclude that the profile theory needs modification regarding (1) crown shape which varies with age and growing space, and (2) the ratio of foliage mass to branch area which varies along the stem. Both aspects emphasise the need to include impacts of disuse of sapwood pipes in models of crown and stem development.     

A protocol for measuring abundance and size of a neotropical liana,Desmoncus polyacanthos (Palmae), in relation to forest structure

This study investigated the relationship between the abundance and size ofD. polyacanthos and measures of forest canopy structure. Várzea and terra firme forest were selected for study at the mouth of the Amazon estuary, Pará, Brazil. The forest canopy characteristics investigated included individual tree height and height to base of the live portion of the crown, forest canopy depth (the distance from the lowest foliage in the understory to the top of the canopy including all free space between top and bottom), forest canopy thickness (the portion of the forest canopy depth occupied only by foliage) and forest canopy density (the sum of lengths of each of the individual crowns that make up forest canopy depth) (see Fig. 2b). The attributes ofD. polyacanthos measured included leaf number, stem height and percent coverage within the stands. Results from this study suggest thatD. polyacanthos can be encouraged to grow through selective cutting of canopy trees and promotion of regenerating forest stands on terra firme fallow. The protocol used to relateD. polyacanthos abundance and size to kind of forest canopy structure can be used for evaluation of other climbing palms.     

Intratree Variation in Fruit Production and Implications for Primate Foraging

We tested the hypothesis that fruit quantity and quality vary vertically within trees. We quantified intratree fruit production before exploitation by frugivores at different heights in 89 trees from 17 species fed on by primates in Kibale National Park, Uganda. We also conducted a pilot study to determine if the nutritional value of fruit varied within tree crowns. Depending on the species and crown size, we divided tree canopies into 2 or 3 vertical layers. In 2-layered trees, upper crowns produced fruits that were 9.6–30.1% bigger and 0.52–140 times the densities of those from lower crowns, with one exception. Among 2-layered trees, upper crowns produced a mean of 46.9 fruits/m³ (median 12.1), while lower crowns produced a mean of 14.1 fruits/m³ (median 2.5). Among 3-layered trees, upper crowns produced a mean density of 49.9 fruits/m³ (median 12.5), middle crowns a mean of 16.8 fruits/m³ (median 6.6), and lower crowns a mean of 12.8 fruits/m³ (median 1.8). Dry pulp and moisture were systematically greater per fruit in the highest compared to the lowest canopy layers (22.4% and 16.4% respectively in 2-layered trees, 49.7% and 21.8% respectively in 3-layered trees). In 1 tree of Diospyros abyssinica, a pilot nutritional study showed that upper crown ripe fruit contained 41.9% more sugar, 8.4% more crude proteins, and 1.8 times less of the potentially toxic saponin than lower crown ripe fruit, but the result needs to be verified with more individuals and species of trees. We discuss the consequences of intratree variations in fruit production with respect to competition among frugivorous primates.