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.     

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.     

Modeling the vertical foliage distribution of an individual <Emphasis Type="Italic">Castanopsis cuspidata</Emphasis> (Thunb.) Schottky,a dominant broad-leaved tree in Japanese warm-temperate forest

The vertical foliage distribution of Castanopsis cuspidata (Thunb.) Schottky was examined in trees of various sizes to clarify its variation in relation to tree size and the light environment in a stand. As indices of these parameters, we analyzed crown social position (CSP: percent of stand height) and specific leaf area (SLA). The vertical foliage distribution of trees was expressed by a Weibull function. The variation in the vertical foliage distribution of C. cuspidata could be categorized into three types using crown social position and light environment. In the first type, leaves were concentrated to the top 20% of the tree; such trees are canopy trees that can receive full sunlight. The second type had a large relative crown depth and an asymmetric distribution with the maximum foliage located near the top of the tree; such trees are suppressed trees whose crowns do not receive sufficient light. The third type had a large relative crown depth and a symmetric distribution; such trees occur in high light environments, although their crowns are in the understory layer. The differences in the vertical foliage distribution are related to the strategies used to capture light. Multiple regression analysis showed that CSP and SLA at the top layer of the tree explained successive changes in the vertical foliage distribution. These results will contribute to scaling-up the vertical foliage distribution to the community level in pure stands of C. cuspidata using an individual-based model.     

Computing competition for light in the GREENLAB model of plant growth: a contribution to the study of the effects of density on resource acquisition and architectural development

BACKGROUND AND AIMS: The dynamical system of plant growth GREENLAB was originally developed for individual plants, without explicitly taking into account interplant competition for light. Inspired by the competition models developed in the context of forest science for mono-specific stands, we propose to adapt the method of crown projection onto the x-y plane to GREENLAB, in order to study the effects of density on resource acquisition and on architectural development. METHODS: The empirical production equation of GREENLAB is extrapolated to stands by computing the exposed photosynthetic foliage area of each plant. The computation is based on the combination of Poisson models of leaf distribution for all the neighbouring plants whose crown projection surfaces overlap. To study the effects of density on architectural development, we link the proposed competition model to the model of interaction between functional growth and structural development introduced by Mathieu (2006, PhD Thesis, Ecole Centrale de Paris, France). KEY RESULTS AND CONCLUSIONS: The model is applied to mono-specific field crops and forest stands. For high-density crops at full cover, the model is shown to be equivalent to the classical equation of field crop production (Howell and Musick, 1985, in Les besoins en eau des cultures; Paris: INRA Editions). However, our method is more accurate at the early stages of growth (before cover) or in the case of intermediate densities. It may potentially account for local effects, such as uneven spacing, variation in the time of plant emergence or variation in seed biomass. The application of the model to trees illustrates the expression of plant plasticity in response to competition for light. Density strongly impacts on tree architectural development through interactions with the source-sink balances during growth. The effects of density on tree height and radial growth that are commonly observed in real stands appear as emerging properties of the model.     

基于不同预测变量的天然椴树可加性地上生物量模型构建

森林生物量是森林生态系统的最基本数量特征,生物量数据是研究许多林业问题和生态问题的基础,因此,准确测定生物量对于计算碳储量以及研究气候变化、森林健康、森林生产力、养分循环等十分重要.目前,测算森林生物量常用的方法为生物量模型估算法.本研究基于小兴安岭地区和张广才岭地区97株实测生物量数据,建立了3个天然椴树立木可加性生物量模型系统(基于胸径的一元可加性生物量模型系统、基于胸径和树高的二元可加性生物量模型系统、基于最优变量的最优可加性生物量模型系统),采用非线性似乎不相关回归法进行参数估计,用加权方法解决模型的异方差问题,并采用“刀切法”进行模型检验.结果表明: 3种可加性生物量模型系统均能较好地对椴树各部分生物量进行拟合和预测(调整后确定系数R_a²>0.84,平均预测误差百分比MPE<8.5%,平均绝对误差MAE<16.3 kg,平均百分标准误差MPSE<28.5%),其中,树干和地上生物量的拟合效果优于树叶、树枝和树冠;在引入树高和树冠因子后,提高了模型的拟合效果和预测能力(R_a²提高0.01～0.04,MAE降低0.01～4.55 kg),缩小了预测值置信区间的范围,树干、树叶和地上生物量提高较多,树枝和树冠提高较少.总体来看,最优生物量模型系统效果最好,其次为二元生物量模型系统,再次是一元生物量模型系统,添加树高和树冠因子进行生物量模型的构建十分必要.     

Understanding changes in black (Picea mariana) and white spruce (Picea glauca) foliage biomass and leaf area characteristics

Key message

Growth conditions related to inter-tree competition greatly influence black and white spruce foliage biomass and projected leaf area characteristics.

Abstract

Foliage characteristics such as biomass and area are important among other reasons because they can be related to tree growth. Despite their economic and ecologic importance, equations to characterize foliage biomass and projected area of black (Picea mariana (Miller) BSP) and white (Picea glauca (Moench) Voss) spruces are sparse. Total foliage biomass and projected leaf area, foliage biomass and leaf area density, and relative vertical distribution of black and white spruces foliage biomass and leaf area were modelled with linear and nonlinear mixed effect models. A total of 65 white spruces and 57 black spruces were destructively sampled at four different locations in Alberta, Québec, and Ontario, Canada. Our results show that for each species, total tree foliage biomass and projected leaf area is proportional to stem diameter, total height, and crown length. The addition of crown length in the equations improved the precision of the predictions of total foliage biomass for both species and diminishes greatly the site level random effect. An increase in DBH for black spruce and in the DBH to total height ratio for white spruce skewed the relative vertical foliage biomass distribution toward the base of the living crown. According to our results, growth conditions or tree development stage influence both foliage biomass and leaf area characteristics of black and white spruces. Our results emphasize the importance of inter-tree competition on foliage biomass characteristics.     

Crown hollowing as a consequence of early shedding of leaves and shoots

Leafing pattern has long been considered as an important element characterizing the growth strategy of tree species; however, the consequences of leafing pattern for tree-crown formation have not been fully understood. To address this issue, the dynamic events (growth, birth, and death) of current-year shoots and leaves were investigated together with their location in saplings of a pioneer tree, Alnus sieboldiana. The leafing pattern was characterized by successive emergence and shedding of short-lived leaves. The combination of successive leafing and within-crown variation in leaf production brought about characteristic outcomes in crown morphology. In the outer crown, because of continuous leaf production, the shoots achieved great extension and enormous daughter shoot production, resulting in rapid expansion of the crown. In contrast, in the inner crown, due to early termination of leaf production, the shoots completely lost their leaves early in the growing season and consequently themselves died and were shed within the season. Such quick shedding of shoots caused “crown hollowing”, i.e., the interior crown consisted of primary branches with little secondary development or foliage. These dynamic features are an effective adaptive strategy in early succession but also may be a disadvantage to maintaining foliage for longer period. Crown maintenance associated with the longevity of structural components is thought to play an important role in survival strategy of tree species.     

樟子松人工林树冠结构的分形分析

基于樟子松人工林7块固定标准地中的31株解析木的树冠体积和叶量,以幂函数关系(F=Av^(D/3))建立了预估树冠表面积的分形维数。同时根据生物量实测数据,建立预估叶量的生物模型Lw=0.180397D^3045903H^-1.67348。基于枝解析、树干解析数据,动态地预估了一年、二年、三年前的树冠体积,并结合树冠体积、叶量的这种幂函数关系可以动态地预估一年、二年、三年前树冠表面积的分形维数,从而反映出树冠结构的动态变化规律。为了了解不同分级样木的分维数变化情况,利用2003年调查的4块生物量标准地数据,根据单株树木各个枝条占据的空间体积与该枝条的带叶枝干重的关系,计算了各标准地不同分级样木树冠的分维数。为探讨单株样木树冠的分维数的计算提供了一种可行方法。树冠的分维数作为表征树冠的动态生长变化是一有用和可靠的指标。     

Light interception in apple trees influenced by canopy architecture manipulation

Improvement of light penetration within tree canopies has been a constant objective of fruit tree architecture manipulation through the setting up of training systems. Recently, centrifugal training, i.e. the removal of fruiting shoots in the tree centre and on the underside of branches, has been proposed to improve fruit size and colour as well as return-bloom as compared to conventional solaxe-trained trees with equivalent crop loads. The present study was conducted to quantify the benefits of centrifugal training on light interception by the fruiting shoots via computer-assisted three-dimensional representations of foliage geometry. Data were collected on six 5-year-old apple trees cv.Galaxy, trained either with solaxe or centrifugal training systems, using an electromagnetic 3D digitiser. The 3D distribution of the foliage in the tree canopy was recreated by combining both the spatial locations of shoots (as measured from 3D digitising) and foliage reconstruction. Light interception efficiency properties of the trees were characterised by silhouette to total area ratio (STAR) values computed from images of the 3D mock-ups. Compared to the solaxe system, centrifugal training significantly improved the STAR of the whole tree by 20%. It also increased both leaf area and STAR of the fruiting shoots by approximately 15%, regardless of their position in the canopy. In this paper, we discuss the role of this enhanced light interception by the canopy in increasing the autonomy of the fruiting shoot, i.e. improved fruit size and colour, and return-bloom.     

Spatial distribution of leaf water-use efficiency and carbon isotope discrimination within an isolated tree crown

The spatial variations in the stable carbon isotope composition (δ¹³C) of air and leaves (total matter and soluble sugars) were quantified within the crown of a well‐watered, 20‐year‐old walnut tree growing in a low‐density orchard. The observed leaf carbon isotope discrimination (Δ) was compared with that computed by a three‐dimensional model simulating the intracanopy distribution of irradiance, transpiration and photosynthesis (previously parameterized and tested for the same tree canopy) coupled to a biophysically based model of carbon isotope discrimination. The importance of discrimination associated with CO₂ gradients encountered from the substomatal sites to the carboxylation sites was evaluated. We also assessed by simulation the effect of current irradiance on leaf gas exchange and the effect of long‐term acclimation of photosynthetic capacity and stomatal and internal conductances to light regime on intracanopy gradients in Δ. The main conclusions of this study are: (i) leaf Δ can exhibit important variations (5 and 8‰ in total leaf material and soluble sugars, respectively) along light gradients within the foliage of an isolated tree; (ii) internal conductance must be taken into account to adequately predict leaf Δ, and (iii) the spatial variations in Δ and water‐use efficiency resulted from the short‐term response of leaf gas exchange to variations in local irradiance and, to a much lesser extent, from the long‐term acclimation of leaf characteristics to the local light regime.     

LIGNUM: A Tree Model Based on Simple Structural Units

The model LIGNUM treats a tree as a collection of a large numberof simple units which correspond to the organs of the tree.The model describes the three dimensional structure of the treecrown and defines the growth in terms of the metabolism takingplace in these units. The activities of physiological processescan be explicitly related to the tree structures in which theyare taking place. The time step is 1 year. The crown of the model tree consists of tree segments, branchingpoints and buds. Each pair of tree segments is separated bya branching point. The buds produce new tree segments, branchingpoints and buds. The tree segments contain wood, bark and foliage.A model tree consisting of simple elements translates convenientlyto a list structure: the computer program implementing LIGNUMtreats the tree as a collection of lists. The annual growth of the tree is driven by available photosyntheticproducts after respiration losses are accounted for. The photosyntheticrate of foliage depends on the amount of light. The amount ofphotosynthates allocated to the growth of new tree segmentsis controlled by the light conditions and the amount of foliageon the mother tree segment. In principle, the biomass relationshipsof the tree parts follow the pipe model hypothesis. The orientationof new tree segments results from the application of constantbranching angles. LIGNUM has been parametrized for young Scotspine (Pinus sylvestrisL.) trees. However, the model is generic;with a change of parameter values and minor modifications itcan be applied to other species as well. Growth model; object-oriented modelling; tree architecture; branching structure; Pinus sylvestrisL.; developmental morphology and physiology; photosynthesis; respiration     

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.     

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.     

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.     

Distributional gradients and variability of macroelement concentrations in the crowns of plantation grownPinus resinosa (AIT.)

Summary The vertical distribution of inorganic nutrient concentrations in red pine were dependent on the foliage age. Older foliage did not show an average vertical gradient while younger foliage did show a significant gradient. Horizontal gradients across foliage age classes in a branch whorl were consistent for all branch whorls, but the relative difference between the concentration of the current foliage and foliage three years-old or older was dependent on crown position. Coefficients of variation (CV) did not show a variability gradient in the crown for nitrogen and phosphorus. Variability of potassium tended to decrease as foliage age increased. Contribution of the University of Florida, Soil Science Department, Gainesville, FL and State University of New York, College of Environmental Science and Forestry, Syracuse, New York. Florida Agricultural Experiment Station Journal Series No. 3017.     

Crown allometries are less responsive than stem allometry to tree size and habitat variations in an Indian monsoon forest

While theoretical allometric models postulate universal scaling exponents, empirical relationships between tree dimensions show marked variability that reflects changes in the biomass allocation pattern. As growth of the various tree compartments may be controlled by different functions, it is hypothesized that they may respond differently to factors of variation, resulting in variable tree morphologies and potentially in trade-offs between allometric relationships. We explore the variability of tree stem and crown allometries using a dataset of 1,729 trees located in an undisturbed wet evergreen forest of the Western Ghats, India. We specifically test whether species adult stature, terrain slope, tree size and crown light exposure affect the relationships between stem diameter and stem height (stem allometry), and between stem diameter and crown width, crown area and crown volume (crown allometries). Results show that both stem and crown allometries are subject to variations in relation to both endogenous (tree size, species adult stature) and exogenous (terrain slope, crown light exposure) factors. Stem allometry appears to be more affected by these factors than are crown allometries, including the stem diameter–crown volume relationship, which proved to be particularly stable. Our results support the idea that height is a prevailing adjustment factor for a tree facing variable growth (notably light) conditions, while stem diameter–crown volume allometry responds more to internal metabolic constraints. We ultimately discuss the various sources of variability in the stem and crown allometries of tropical trees that likely play an important role in forest community dynamics.     

Canopy structures and its effect on shoot growth and flowering in subalpine forests

Changes in foliage density distribution with altitude and its effect on shoot growth and flowering were studied in forest section planes (profiles) of subalpine forests and scrubs (Krummholz) in Nepal and Japan.Patterns of foliage in forest canopy were evaluated by an analysis of variance. Foliage densities were very high at high altitude canopies, but the change in spatial patterns along altitude differs in both areas.The spatial pattern of new shoot production was similar to that of current foliage density and was affected by the amount of foliage above the sample probably through light condition. Flowering of tall trees occurred in the surface of the upper canopy, but a shrub species flowered even under tree canopies.     

Simple equations to estimate light interception by isolated trees from canopy structure features: assessment with three-dimensional digitized apple trees

* Simple models of light interception are useful to identify the key structural parameters involved in light capture. We developed such models for isolated trees and tested them with virtual experiments. Light interception was decomposed into the projection of the crown envelope and the crown porosity. The latter was related to tree structure parameters. * Virtual experiments were conducted with three-dimensional (3-D) digitized apple trees grown in Lebanon and Switzerland, with different cultivars and training. The digitized trees allowed actual values of canopy structure (total leaf area, crown volume, foliage inclination angle, variance of leaf area density) and light interception properties (projected leaf area, silhouette to total area ratio, porosity, dispersion parameters) to be computed, and relationships between structure and interception variables to be derived. * The projected envelope area was related to crown volume with a power function of exponent 2/3. Crown porosity was a negative exponential function of mean optical density, that is, the ratio between total leaf area and the projected envelope area. The leaf dispersion parameter was a negative linear function of the relative variance of leaf area density in the crown volume. * The resulting models were expressed as two single equations. After calibration, model outputs were very close to values computed from the 3-D digitized databases.     

Crown asymmetry in high latitude forests: disentangling the directional effects of tree competition and solar radiation

Light foraging by trees is a fundamental process shaping forest communities. In heterogeneous light environments this behavior is expressed as plasticity of tree growth and the development of structural asymmetries. We studied the relative influence of neighborhood structure and directional solar radiation on horizontal asymmetry of tree crowns in late‐successional high latitude (67–68°N) forests in northern Fennoscandia. We described crown asymmetries as crown vectors (i.e. horizontal vectors from stem center to crown center), which we obtained from canopy maps based on crown perimeter measurements in the field. To disentangle the influence of the two main determinants, inter‐tree competition and directionality of above‐canopy solar radiation at high latitudes, we applied circular statistical models, utilizing cylindrical distributions, to these data consisting of orientations and intensities of crown asymmetry. At the individual tree level, our model predicted crown asymmetry vectors from the current stand structure, and the predictions became better when the intensity of asymmetry (i.e. crown vector length) was higher. Competition was the main determinant of crown asymmetry for 2/3 of trees, and the model predictions improved when we incorporated the directionality of solar radiation. At the stand‐level, these asymmetries had resulted in a small increment of the projected canopy area and an increased regularity of spatial structure. Our circular statistical modelling approach provided a quantitative evaluation of the relative importance of directionality of solar radiation and neighborhood stand structure, showing how both of these factors play a role in formation of crown asymmetries in high latitude forests. This approach further demonstrated the applicability of circular statistical modeling in ecological studies where the response variable has both orientation and intensity.     

Ground monitoring the light–shadow windows of a tree canopy to yield canopy light interception and morphological traits

Monitoring the light–shadow windows of a tree via a grid system on the ground was performed on sunny summer days at high spatial resolution using a custom‐built, inexpensive scanner. The measurements were taken with two goals: (1) to quickly and remotely quantify the overall, short‐wave solar radiation (300–1100 nm) intercepted by the tree canopy, and (2) to yield such crown geometric traits as shape, size and the number of theoretical canopy leaf layers (leaf layer index, LLI) in relation to the section orthogonal to sunbeam direction (sun window). The ground readings at each measurement over the day were used to project a digitized shadow image. Image processing was applied and the intercepted radiation was calculated as the difference from the corresponding incoming radiation above the canopy. Tree‐crown size and shape were profiled via computer imaging by analysing the different shadow images acquired at the various solar positions during the day. It is notable that these combined images yielded the crown features without having to parameterize such canopy characteristics as foliage extension and spatial distribution.