北京山区干旱胁迫下侧柏叶片水分吸收策略

干旱与半干旱地区,水分是限制树木生长的重要影响因子。由于降水稀缺且分配不均,叶片吸收水分是此地区树木吸收和利用小量级降水和凝结水的主要方式。北京山区处于易旱少雨的生态脆弱地带,森林植被经常遭受干旱胁迫,所以对该地区的森林系统而言,叶片直接吸收利用截留的降雨是干旱时期树木获得水分的重要途径。基于野外对比控制试验和室内盆栽模拟试验,选取北京山区的主要造林树种侧柏为研究对象,进行利用天然降雨与模拟降雨试验,研究降雨前后侧柏叶片吸水特征,探究侧柏在干旱环境下如何通过叶片吸水缓解干旱胁迫。结果表明：当侧柏长期处于干旱胁迫状态时,叶片可以利用降雨,从中获益用来缓解树木的干旱胁迫状态;叶片的吸水能力与降雨强度呈正相关关系,与土壤含水率呈负相关关系;重度干旱下侧柏植株在降雨强度为15 mm/h时叶片吸水现象最明显,叶水势变化最大为(1.18±0.17) MPa,叶片含水率变化最大为(8.47±1.00)mg/cm~2;当土壤水率高于20.8%时,基本不发生叶片吸水现象。试验结果说明在干旱地区叶片吸水是树木除根系吸水外的重要水分来源方式,并且对干旱地区有效利用短缺水资源,减轻植物水分亏缺具有重要意义。     

东亚地区夏季干旱、强台风事件与松树枯萎病的关系

松树枯萎病的发生和流行给东亚的日本、中国和韩国带来了不小的损失,引起许多国家的重视. 本文应用实地观测和气象数据分析等方法,研究了东亚地区松树枯萎病的发生和流行特点,以及与灾害气象事件之间的关系.结果表明: 在日本、中国和韩国,持续的夏季干旱少雨和强台风等极端气象事件能够诱发松树枯萎.在极端干热的环境中,松树常出现能量代谢失调, 以至于受胁迫的松树整株枯萎;而在低温多雨的年份,松树枯萎少,甚至没有发生枯死现象.在松材线虫及其媒介昆虫侵染之前,松树的活力业已下降.松树枯萎病似乎应该局限在台风频发和持续干热的地区.在自然环境优越、少有台风和干热事件出现且没有不当扩大松树栽培范围的地区,松树枯萎病大面积发生的可能性不大.     

Acclimation of crown structure to drought in Quercus robur L.—intra- and inter-annual variation of abscission and traits of shed twigs

In pedunculate oak (Quercus robur L.) the architecture of the crown is strongly influenced by age and vigour of the tree. In old oak trees cladoptosis is a major mechanism on the transformation of crown architecture. Although it can be seen quite regularly, the causes and timing of shedding of twigs as well as the quantity of affected branches remain unclear. Because abscission is often used as an indicator of reduced vigour or stress in the assessment of stand and ornamental tree health, it deserves detailed investigation, especially in the context of oak decline. We studied the inter- and intra-annual variation of abscission in six stands across the eastern part of Germany in order to identify possible triggering events and controlling factors of abscission.On average, the number of twigs abscised per year increased from 1999 to 2001. While in 1999 approximately 100 abscised twigs per m² per year were shed, this number increased to 250 per m² in 2001. The majority of twigs was actively shed, a significant proportion of the remaining twigs was partly abscised. Only a small part of the abscised twigs had leaves attached to them. From June to September 20 per cent of the twigs had leaves, in the remaining months of the year less than 10 per cent.The analysis of almost 30,000 twig fragments over the course of 3 years demonstrated at least partial control by the tree of the process of abscission. The loss of the terminal bud and the formation of a male flower are traits correlated with abscission.Our data do not support the widely held belief that cladoptosis is mainly an immediate reaction to drought stress that reduces transpiring leaf area. The proportion of leafy twigs was quite low and the main peak of abscission occurred in late autumn, thus having little immediate effect on transpiration. In addition, we observed a time lag of 3 weeks between the onset of drought stress and increased levels of abscission.     

Variation in litter under individual tree crowns: Implications for scattered tree ecosystems

In forest ecosystems litter is usually assessed in terms of the average amount produced by the canopy. In scattered tree ecosystems this approach is problematic because the canopy is discontinuous and the spatial arrangement of litter highly variable. We addressed this problem by quantifying the spatial variation in litter load and litter composition associated with individual trees in a Eucalyptus melliodora – Eucalyptus blakelyi woodland. Litter was sampled under crowns and in grassland adjacent to 10 E. blakelyi and 10 E. melliodora trees ranging in diameter at breast height (dbh) from 14 to 129 cm. A total of 302 L samples were collected from these trees, at distances ranging from 0 to 42 m from main stem. The sampled litter loads ranged from 0.02 to 109.3 t ha^?1 and were significantly higher under tree crowns than in grassland for litter and each component of litter (leaves, bark, fine twigs, coarse twigs). In particular, the mean litter load under tree crowns (12.5 t ha^?1) was an order of magnitude higher than the mean litter load in grassland (1.27 t ha^?1). There was a significant (P = 0.0103) positive relationship between mean litter load under the tree crown and dbh, indicating larger trees produced more litter per unit area of ground than smaller trees. Generalized Linear Modelling produced highly significant (P < 0.0001) models predicting the spatial variation in litter load and litter composition in terms of distance from main stem and dbh. Our models demonstrate gradients in litter load and composition under tree crowns. These gradients were most pronounced for the large trees in our study. The disproportionate input of litter and variety of litter components associated with large trees in our study supports their keystone role in scattered tree ecosystems and highlights the need to maintain these structures in agricultural landscapes.     

河北太行山南段树木年轮指示的167年来相对湿度变化研究

研究建立了河北太行山南段侧柏(Platycladus orientalis)树轮宽度年表,分析树木径向生长对水热变化的响应,构建了研究区167年来相对湿度变化序列。结果表明,侧柏树轮宽度与6月降水显著正相关,与5—7月逐月均温和最高温均显著负相关,侧柏径向生长受到春末夏初水热条件的显著制约。研究区167年来干湿变化明显,明显的干旱期有1876—1877,1900—1901,1904—1912,1918—1921,1926—1930和1933—1935年,湿润期有1871—1873,1882—1884,1888—1890,1893—1895,1953—1956,1971—1972和2002—2003年。5—7月相对湿度变化与旱涝等级负相关达到0.01显著水平。周期分析表明,167年来5—7月相对湿度变化存在2—4年、7.71年和60年左右周期,可能与太平洋活动、太阳活动有一定关联。     

Seeing the trees for the leaves – oaks as mosaics for a host-specific moth

From the perspective of a specialist herbivore, how homogenous are individual tree crowns as patches of habitat? We partitioned variation in physical and chemical host leaf traits and in the abundance and performance of a specialist oak leaf miner, Tischeria ekebladella, into variation at different hierarchical levels. For the phenolic contents of the leaves, we examined variation among oak stands, among trees within stands and among branches within trees. For leaf size and water content, we assessed variation among trees within a single stand, among shoots within trees, and among leaves within shoots. For moth abundance and performance, we examined variation across all levels: among oak stands, among trees within stands, among branches within trees, among shoots within branches and among leaves and insect individuals. For measures of phenolic contents, we found little variation among stands but substantial variation among individual trees. Yet, a tree particularly rich in a given compound was often comparatively poor in another. At a finer spatial scale, the phenolic composition of individual parts of a single tree was quite consistent, whereas leaf weight and water content varied widely within individual tree crowns. Moth abundances varied more among shoots within branches than at any other spatial level, whereas moth survival showed equal levels of variation within individual shoots as among separate oak stands. Likewise, for four other measures of larval performance (assessed at the level of trees and lower), we found more variation within than between trees. In conclusion, the large variation observed in the performance of a specialist moth and in the physical traits of the leaves among different parts of single tree crowns refutes the image of an oak tree as an ‘island’ of internally homogeneous quality. Hence, we may expect little evolutionary adaptation of T. ekebladella at the scale of individual trees. The moths may instead evolve to behaviourally select their resource at a very fine scale. Large variation within trees also calls for extensive replication within trees in ecological sampling designs and/or the sampling of maximally similar leaves.     

江西省不同类型退化荒山生态系统植被恢复与重建措施

对江西省 5种不同类型退化荒山生态系统进行综合治理 ,开展植被恢复与重建技术及理论研究。研究结果表明 :优选的植物组合及合理的配置并辅以工程措施是快速启动植被恢复进程 ,控制水土流失、重建退化荒山生态系统的有效措施。一林多用的树种组合 ,合理的生物体系设计和针、阔叶树种混交是改良地力 ,改善环境 ,促进退化荒山生态系统进展演替的优良途径。先进造林技术的应用和林农牧业结合的复合经营方式是提高综合治理效益 ,促进并维持生态恢复过程稳定的先决条件     

Evidence that branch cuvettes are reasonable surrogates for estimating O3 effects in entire tree crowns

Within the scope of quantifying ozone (O(3)) effects on forest tree crowns it is still an open question whether cuvette branches of adult trees are reasonable surrogates for O(3) responses of entire tree crowns and whether twigs exhibit autonomy in defense metabolism in addition to carbon autonomy. Therefore, cuvette-enclosed branches of mature beech (Fagus sylvatica) trees were compared with branches exposed to the same and different ozone regimes by a free-air fumigation system under natural stand conditions by means of a VICE VERSA experiment. For this purpose, cuvettes receiving 1 x O(3) air were mounted in trees exposed to 2 x O(3) and cuvettes receiving 2 x O(3) air were mounted in trees exposed to 1 x O (3) in the upper sun crown. At the end of the fumigation period in September 2004, leaves were examined for differences in gas exchange parameters, pigments, antioxidants, carbohydrates, and stable isotope ratios. No significant differences in foliar gas exchange, total carbohydrates, stable isotope ratios, pigment, and antioxidant contents were found as a consequence of cuvette enclosure (cuvette versus free-air branches) of the same O(3) concentrations besides increase of glucose inside the cuvettes and reduction of the de-epoxidation state of the xanthophyll cycle pigments. No significant ozone effect was found for the investigated gas exchange and most biochemical parameters. The total and oxidized glutathione level of the leaves was increased by the 2 x O(3) treatment in the cuvette and the free-air branches, but this effect was significant only for the free-air branches. From these results we conclude that cuvette branches are useful surrogates for examining the response of entire tree crowns to elevated O(3) and that the defence metabolism of twigs seems to be at least partially autonomous.     

Growth and resilience responses of Scots pine to extreme droughts across Europe depend on predrought growth conditions

Global climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees’ phenotypic variability, which is, in turn, affected by long‐term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree‐level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree‐, site‐, and drought‐related factors and their interactions driving the tree‐level resilience to extreme droughts. We used a tree‐ring network of the widely distributed Scots pine (Pinus sylvestris) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long‐term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation.     

Drought and cold spells trigger dieback of temperate oak and beech forests in northern Spain

Dieback in temperate forests is understudied, despite this biome is predicted to be increasingly affected by more extreme climate events in a warmer world. To evaluate the potential drivers of dieback we reconstructed changes in radial growth and intrinsic water-use efficiency (iWUE) from stable isotopes in tree rings. Particularly, we compared tree size, radial-growth trends, growth responses to climate (temperature, precipitation, cloudiness, number of foggy days) and drought, and changes in iWUE of declining and non-declining trees showing contrasting canopy dieback and defoliation. This comparison was done in six temperate forests located in northern Spain and based on three broadleaved tree species (Quercus robur, Quercus humilis, Fagus sylvatica). Declining trees presented lower radial-growth rates than their non-declining counterparts and tended to show lower growth variability, but not in all sites. The growth divergence between declining and non-declining trees was significant and lasted more in Q. robur (15–30 years) than in F. sylvatica (5–10 years) sites. Dieback was linked to summer drought and associated atmospheric patterns, but in the wettest Q. robur sites cold spells contributed to the growth decline. In contrast, F. sylvatica was the species most responsive to summer drought in terms of growth reduction followed by Q. humilis which showed coupled changes in growth and iWUE as a function of tree vigour. Low growth rates and higher iWUE characterized declining Q. robur and F. sylvatica trees. However, declining F. sylvatica trees became less water-use efficient close to the dieback onset, which could indicate impending tree death. In temperate forests, dieback and growth decline can be triggered by climate extremes such as dry and cold spells, and amplified by climate warming and rising drought stress.     

Balancing the risks of hydraulic failure and carbon starvation: a twig scale analysis in declining Scots pine

Understanding physiological processes involved in drought‐induced mortality is important for predicting the future of forests and for modelling the carbon and water cycles. Recent research has highlighted the variable risks of carbon starvation and hydraulic failure in drought‐exposed trees. However, little is known about the specific responses of leaves and supporting twigs, despite their critical role in balancing carbon acquisition and water loss. Comparing healthy (non‐defoliated) and unhealthy (defoliated) Scots pine at the same site, we measured the physiological variables involved in regulating carbon and water resources. Defoliated trees showed different responses to summer drought compared with non‐defoliated trees. Defoliated trees maintained gas exchange while non‐defoliated trees reduced photosynthesis and transpiration during the drought period. At the branch scale, very few differences were observed in non‐structural carbohydrate concentrations between health classes. However, defoliated trees tended to have lower water potentials and smaller hydraulic safety margins. While non‐defoliated trees showed a typical response to drought for an isohydric species, the physiology appears to be driven in defoliated trees by the need to maintain carbon resources in twigs. These responses put defoliated trees at higher risk of branch hydraulic failure and help explain the interaction between carbon starvation and hydraulic failure in dying trees.     

根系功能性状对干瘠立地适应的种间差异——以北京石质山地主要观赏树种为例

分析干瘠立地中不同树种根功能性状的变化及差异,研究不同树种对干瘠环境的适应对策。试验材料为林龄、林分密度相对一致的石质山区主要观赏树种（黄栌、山桃、栓皮栎）,采用挖掘法获取树木细根,测定细根形态参数。根据田间持水量和石砾含量分别设置4个干瘠梯度（轻度、中度、重度、极度）,对比分析北京干瘠地3种树种根功能性状,探讨干瘠立地中不同树种根系形态的差异及其对特殊生境的适应。结果表明：干旱、瘠薄均对树木的根功能性状产生影响,但对不同性状以及不同树种的影响程度具有差异,其中对山桃根系的影响最大;3种树种根功能性状在不同干瘠等级下存在显著变化,同一性状的变化因树种不同而异。从轻度到极度干旱,3种树种的RL均逐渐减小,但是黄栌变化不显著;黄栌、栓皮栎RTD随干瘠的加剧而减小,而山桃随干瘠的加剧而增加;黄栌和山桃SRL随干旱、瘠薄的加剧,表现出一致的规律,即从轻度到极度,其SRL随即减小,而栓皮栎SRL随干旱、瘠薄的加剧而增大;随干瘠的加剧,黄栌SRA显著减小,山桃、栓皮栎SRA随干瘠的加剧而增大。树木对干瘠环境的变化都有一定的响应能力及适应对策,能够通过自身的形态变化来适应不断变化的环境,但不同物种对干瘠环境变化的适应途径不同,通过权衡碳分配、碳消耗采取了相应的生态适应策略,不同树种间存在明显差异。本研究3种树种中,黄栌在干瘠环境下根系性状变化不显著,耐干瘠能力较强;干瘠胁迫对山桃根系性状影响显著,山桃通过主动调节根系形态变化来适应不断变化的环境,具有较强的抗干瘠能力;栓皮栎的细根数量多,吸收土壤水分、养分能力较强,能够在干瘠环境中快速生长。     

Patterns of Foraging of the Forest Ant Formica neogagates Emery (Hymenoptera: Formicidae) on Tree Branches

Numbers of ants moving on trunks of oak trees and successive visits of workers of the forest ant Formica neogagates Emery to leaves of black birch trees were recorded in the forest. Ants were found to search systematically because they tended to move in the same direction on a twig before and after visiting a leaf. However, many leaves were skipped, and the probability of visiting an encountered leaf was about 0.5. Data were used to develop a computer model of ant foraging on leaves and twigs of trees. It was found that reasonable model outputs of numbers of leaves skipped between visits and the number of new leaves visited, as the total number of visits increased, could be obtained only if ants were assumed to partially avoid leaves that they had previously visited. Model results implied that 100 ants in a tree foraging for about 3 h should be able to visit about one quarter of the leaves in a tree.     

Spatio-temporal dynamics of endophyte diversity in the canopy of European ash (Fraxinus excelsior)

Leaf-inhabiting endophytic fungi of Fraxinus excelsior growing in a floodplain forest were isolated during 2008 to investigate vertical community structure, species richness and seasonal variation. The analysis of 848 fungal endophytes from 213 leaves resulted in 50 different species. In the understorey, infection density and species richness were higher than in the crowns of mature trees throughout the whole vegetation period. Within tree crowns, sun-exposed leaves of the top canopy exhibited the lowest infection rates. Most species were rare or absent in spring and in the light crowns and frequent in autumn and the understorey. However, some species, especially the two most frequent, Alternaria infectoria and A. alternata, deviated from these patterns. Young leaves were nearly free of endophytes. Apparently, the subsequent infection and establishment of fungi strongly depend on microclimatic parameters and leaf characters, which create highly variable spatial and temporal colonisation patterns within an individual tree.     

Image measurements of leaf scorches on landscape trees subjected to extreme meteorological events

Trees grow in changing environments and are usually injured by many external factors including both abiotic and biotic. Leaf scorch is a main symptom for many landscape trees when they are stricken by meteorological extreme events, such as summer drought, strong typhoon and winter freezing and so on. In the study, the internal angle of injured area (IAIA) was defined in relation to vein lines and measured non-destructively for some landscape tree species. Meanwhile, the leaf scorch area percent (LSAP) and percent of scorched central vein (PSCV) were also determined from RGB images. Positive relationships between IAIA and LSAP and between PSCV and LSAP were found. The larger IAIA values for scorched leaves and smaller IAIA values for spot/anthracnose diseased leaves of the studied trees indicated that IAIA may be used as a diagnosing index of leaf scorch symptoms.     

干旱对兴安落叶松枝叶非结构性碳水化合物的影响

降水格局的变化以及极端干旱的频繁发生是全球气候变化的重要特征之一。为了揭示干旱对树木碳代谢的影响,通过控雨试验研究兴安落叶松(Larix gmelinii Rupr.)枝叶的非结构性碳水化合物(NSC)及其组分(可溶性糖和淀粉)的浓度对降水减少的响应,探索枝叶NSC浓度与土壤含水率的关系。控雨试验包括减雨100%(100%RE)、减雨50%(50%RE)和对照(CK)3个处理;控雨时期为2012年生长季(6月至8月)。结果表明,叶NSC浓度对干旱处理的响应比枝更显著。控雨处理对枝叶总NSC浓度影响不显著(P0.05),试验期间叶总NSC平均浓度变化在9.45—14.12 mg/g范围内;枝总NSC平均浓度变化在7.72—9.26 mg/g之间。然而,不同处理之间的叶片可溶性糖浓度差异显著。100%RE最高(8.98±0.31)mg/g、50%RE次之(8.45±0.13)mg/g、CK最低(7.73±0.32)mg/g。相反,叶淀粉浓度以CK最高(2.99±0.22)mg/g、50%RE次之(2.68±0.32)mg/g、100%RE最低(2.63±0.17)mg/g。叶可溶性糖与淀粉浓度的比值的大小顺序为:CK(2.27)50%RE(2.51)100%RE(3.70)。叶可溶性糖浓度、可溶性糖浓度和淀粉浓度的比值与土壤含水率呈显著的负相关关系(P0.05),而叶淀粉浓度有随土壤含水率升高而增高趋势,但相关关系不显著(P0.05)。叶NSC总浓度、枝NSC及其组分浓度与土壤含水率的关系均不显著(P0.05)。研究表明,短期干旱对兴安落叶松树体内总NSC浓度的影响不显著,树木可以通过将淀粉转化成可溶性糖的方式维持其正常的呼吸作用等生理活动。     

Leaf dimorphism in Thuja plicata and Platycladus orientalis (thujoid Cupressaceae s. str., Coniferales): the changes in morphology and anatomy from juvenile needle leaves to mature scale leaves

Thuja plicata and Platycladus orientalis initially produce only bifacial needle leaves. When the first lateral shoots develop, the leaf morphology and anatomy changes dramatically. Subsequently, only greatly reduced, bifacial scale leaves are developed. A new kind of “superimposed bifaciality” occurs with the change from juvenile needle leaves to mature scale leaves. Anatomical dorsiventrality affects not only the individual leaf, but also the complete plagiotropic lateral shoots of Thuja, which have a sun- and shade-exposed side. The upper light-exposed median leaves show adaxial leaf anatomy, contrary to the lower shaded median leaves showing abaxial leaf anatomy. Due to their mixed exposure, the lateral leaves show a lateral differentiation. At vertical lateral shoots of Platycladus, a predominant light-exposed side is absent. Thus, the anatomical dorsiventrality does not affect the complete shoot. Here the morphological abaxial side of a scale leaf becomes functionally and physiologically adaxial by reorientation of the palisade parenchyma and stomata. In juvenile needle leaves, the palisade parenchyma is located adaxial, with the majority of stomata being located abaxial. Conversely, in mature scale leaves, the palisade parenchyma is abaxial and the majority of stomata are adaxial.     

Winter drying injury of Ligustrum lucidum associated with water and energy imbalance

To prove the theory that meteorological factors, especially winter drying events, usually affect trees by inducing water and energy imbalance, systematic studies of winter drying damage to Ligustrum lucidum trees in Shandong, China in 2018 were performed by analyzing digital and thermal images of the trees and integrating the results with meteorological and geostatistical analysis and the results of osmotic stress tests in saturated sucrose solution. The results indicate that sap water can not only be transported upward with assistance from cohesion-tension force, but also can be turned back from the terminals of leaves and branches under the inverse pulling force. Under extreme conditions during winter drying events, the breakage of sap water column and reduction of the terminal tissues or organs occur. Similar “∧”-shaped leaf symptoms can also be induced by freezing, summer drought and osmotic stress. The results support the theory of transpiration surface reduction characterized by unity of freezing and drying. Therefore, severe damage to juvenile plants of Ligustrum lucidum under these conditions may be caused by the combined effects of low temperature, drought, strong irradiation and saline soil as well as by the sensitive status of the plants.     

The expression of light-related leaf functional traits depends on the location of individual leaves within the crown of isolated Olea europaea trees

Background The spatial arrangement and expression of foliar syndromes within tree crowns can reflect the coupling between crown form and function in a given environment. Isolated trees subjected to high irradiance and concomitant stress may adjust leaf phenotypes to cope with environmental gradients that are heterogeneous in space and time within the tree crown. The distinct expression of leaf phenotypes among crown positions could lead to complementary patterns in light interception at the crown scale.Methods We quantified eight light-related leaf traits across 12 crown positions of ten isolated Olea europaea trees in the field. Specifically, we investigated whether the phenotypic expression of foliar traits differed among crown sectors and layers and five periods of the day from sunrise to sunset. We investigated the consequences in terms of the exposed area of the leaves at the tree scale during a single day.Key Results All traits differed among crown positions except the length-to-width ratio of the leaves. We found a strong complementarity in the patterns of the potential exposed area of the leaves among day periods as a result of a non-random distribution of leaf angles across the crown. Leaf exposure at the outer layer was below 60 % of the displayed surface, reaching maximum interception during morning periods. Daily interception increased towards the inner layer, achieving consecutive maximization from east to west positions within the crown, matching the sun’s trajectory.Conclusions The expression of leaf traits within isolated trees of O. europaea varies continuously through the crown in a gradient of leaf morphotypes and leaf angles depending on the exposure and location of individual leaves. The distribution of light-related traits within the crown and the complementarity in the potential exposure patterns of the leaves during the day challenges the assumption of low trait variability within individuals.