Plant growth-form alters the relationship between foliar morphology and species shade-tolerance ranking in temperate woody taxa

Variation in leaf size (area per leaf) and leaf dry weight per area (LWA) in relation to species shade- and drought-tolerance, characterised by Ellenberg's light (ELD) and water demand (EWD) values, respectively, were examined in 60 temperate woody taxa at constant relative irradiance. LWA was independent of plant size, but leaf size increased with total plant height at constant ELD. Canopy position also affected leaf morphology: leaves from the upper crown third had higher LWA and were larger than leaves from the lower third. Leaf size and LWA were negatively correlated, and leaf size decreased and LWA increased with decreasing species shade-tolerance. Mean LWA was similar for trees and shrubs, but trees had larger leaves than shrubs. Furthermore, all relationships were altered by plant growth-form: none of the qualitative tendencies was significant for trees. This implies the considerably lower plasticity of foliar parameters in trees than those in shrubs. Accordingly, shade-tolerance of trees, having relatively constant leaf structure, may be most affected by the variability in biomass partitioning and crown geometry which influence foliage distribution and spacing and finally determine canopy light absorptance. Alteration of leaf form and investment pattern for construction of unit foliar surface area which change the efficiency of light interception per unit biomass investment in leaves, is a competitive strategy inherent to shrubs. EWD as well as wood anatomy did not control LWA and leaf size, though there was a trend of ring-porous tree species to be more shade-tolerant than diffuse-porous trees. Since ring-porous species are more vulnerable to cavitation than diffuse-porous species, they may be constrained to environments where irradiances and consequently evaporative demand is lower.     

Energy requirement for foliage formation is not constant along canopy light gradients in temperate deciduous trees

Foliage construction cost (glucose requirement for formation of a unit foliar biomass, G , kg glu kg⁻¹), chemical composition and morphology were examined along a light gradient across the canopies in five deciduous species, which ranked according to increasing shade-tolerance as Populus tremula < Fraxinus excelsior < Tilia cordata = Corylus avellana < Fagus sylvatica . Light conditions in the canopy were estimated by a hemispheric photographic technique, allowing ranking of sample locations according to long-term light input incident to the sampled leaves (relative irradiance). G and foliage carbon concentration increased with increasing relative irradiance in F. excelsior , T. cordata and C. avellana , but wereindependent of irradiance in F. sylvatica and P. tremula . However, if G of non-structural-carbohydrate-free dry mass was considered, it also increased with increasing relative irradiance in P. tremula . A positive correlation between the concentration of carbon-rich lignin and irradiance, probably a result of the acclimation to greater water stress at higher light, was the major reason for the light-dependence of G . Lignin concentrations were highest in more shade-tolerant species, resulting in greatest carbon concentrations in these species. Since carbon concentration and G are directly linked, the leaves of shade-tolerant species were also more expensive to construct. As the result of these effects, G increased faster with increasing leaf dry mass per area which was mainly determined by relative irradiance, in shade-tolerators. Given that shade-tolerant species had lower leaf dry mass per area at common irradiance and that this saturated at lower relative irradiance than leaf dry mass per area in the intolerant species, it was concluded that enhanced energy requirements for foliage construction might constrain species morphological plasticity and the upper limit of leaf dry mass per area attainable at high light.     

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.     

Is the Lower Shade Tolerance of Scots Pine,Relative to Pedunculate Oak,Related to the Composition of Photosynthetic Pigments?

Foliage of Scots pine (Pinus sylvestris L.) and pedunculate oak (Quercus robur L.) was collected in a mixed pine/oak forest at canopy positions differing in radiation environment. In both species, chlorophyll (Chl) a/b ratios were higher in foliage of canopy positions exposed to higher irradiance as compared to more shaded crown layers. Throughout the growing season, pine needles exhibited significantly lower Chl a/b ratios than oak leaves acclimated to a similar photon availability. Hence, pine needles showed shade-type pigment characteristics relative to foliage of oak. At a given radiation environment, pine needles tended to contain more neoxanthin and lutein per unit of Chl than oak leaves. The differences in pigment composition between foliage of pine and oak can be explained by a higher ratio of outer antennae Chl to core complex Chl in needles of P. sylvestris which enhances the efficiency of photon capture under limiting irradiance. The shade-type pigment composition of pine relative to oak foliage could have been due to a reduced mesophyll internal photon exposure of chloroplasts in needles of Scots pine, resulting from their xeromorphic anatomy. Hence, the higher drought tolerance of pine needles could be achieved at the expense of shade tolerance.     

Leaf Biomechanics and Biomass Investment in Support in Relation to Long-Term Irradiance in Fagus

Abstract: We investigated biomass investment in support and assimilative leaf biomass in Fagus orientalis Lipsky and F. sylvatica L., and foliar biomechanical characteristics in F. orientalis to gain mechanistic insight into the determinants of leaf inclination in Fagus along the canopy light gradient. Because the leaf laminas of Fagus are elliptical, with petioles comprising only ca. 8 % of total leaf length, a leaf was approximated as a continuous sine load. Lamina load increased with increasing seasonal integrated quantum flux density in the canopy (Q_int), but leaf length was independent of irradiance. Despite greater load, leaf deflection under leaf own weight was lower for leaves at higher Q_int, indicating that foliage flexural stiffness (EI), that is a variable characterizing the resistance of beam‐like structures to bending, scaled positively with irradiance. The components of EI ‐ the leaf apparent Young's modulus of elasticity (E), which is a measure of leaf material properties, and lamina second moment of area (I), which characterizes the distribution of mass around the axis of bending ‐ were also related to irradiance, with E decreasing, but I increasing with Q_int. The positive scaling of I with Q_int was associated with increases in leaf thickness and, in particular, with increases in the degree of leaf rolling, allowing the distribution of leaf mass further away from the neutral axis. Decreases in E were correlated with decreased leaf biomass investments in the midrib at higher irradiance. Both lamina and midrib nitrogen concentrations decreased with increasing Q_int, suggesting that foliage dry mass based physiological activity was lower at higher irradiance, possibly because of an interaction of Q_int with water stress in the canopy. Given that the veins also provide a pathway for water and nutrient transport to the leaf cells, as well as for carbon translocation from the leaf, lower leaf physiological activity in high light may provide an explanation for the lower biomass investment in major veins in high light. We conclude that foliage biomechanical characteristics and leaf inclination in the canopy are significantly affected by irradiance, and that the light effects may be modified by the reverse correlation between light and water availabilities in the canopy.     

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.     

Inferring changes in foliar mass and area from foliage cover: A mechanistic model

Measuring foliar area or mass directly is destructive, and precludes long‐term, repeated observations of individual trees as they suffer or recover from foliar damage. Instead, foliage cover indices are often used as a proxy for foliar mass. Patterns of fluctuations in foliage cover indices can be used to infer qualitative changes in canopy health. However, foliage cover is not necessarily linearly related to foliar area or mass, and this may confound the detection of significant foliar damage, and comparisons of herbivore browse impacts between individual trees, tree species or sites. I derived a mechanistic model to quantify the relationship between foliar area or mass and foliage cover measured as the proportion of sky occluded by leaves. This one parameter model is close to linear for single‐tiered trees, but increasingly non‐linear for multi‐tiered trees. I compared the non‐linear model to a linear model using foliage cover data from an artificial defoliation experiment on two single‐tiered, sub‐canopy species and from simulated photographic images of single‐ and multi‐tiered canopies. The non‐linear model had lower errors than the linear model, and errors did not increase with foliage density (leaf area per unit area), variation (of leaf sizes within and between canopies) or leaf geometry. The non‐linear model can be easily parameterized from relatively low‐cost observations of foliage cover, independently of empirical measurements of foliar area or mass, and is applicable to a wide range of tree species. It should therefore help managers quantify how changes in foliage cover due to natural fluctuations or foliar damage affect foliar area and mass, and can be used to quantify parameters for models of browse impacts in mixed forest.     

Japanese oak silkmoth feeding preference for and performance on upper-crown and lower-crown leaves

We quantified differences in leaf traits between upper and lower crowns of a deciduous oak, Quercus acutissima, and examined feeding preference, consumption and performance of the Japanese oak silkmoth, Antheraea yamamai, for those leaves. Upper‐crown leaves had significantly smaller area, larger dry mass per area, greater thickness, lower water content, higher nitrogen content and a higher N/C ratio than lower‐crown leaves. When simultaneously offered upper‐crown and lower‐crown leaves, moth larvae consumed a significantly larger amount of the former. However, when fed with either upper‐crown or lower‐crown leaves (no choice), they consumed a significantly larger amount of the latter. Female larvae reared on upper‐crown leaves had a significantly smaller fresh weight, but attained a significantly larger pupal fresh and dry weight, with a significantly higher relative growth rate than those on lower‐crown leaves. Although, like female larvae, male larvae had a significantly smaller fresh weight when reared on upper‐crown leaves, they had a significantly larger value only for pupal dry weight. These results suggest that: (i) larvae ingest a greater amount of lower‐crown leaves to compensate for the lower nitrogen content of the foliage, resulting in having an excess of water because of the higher water content of the foliage; (ii) feeding preference for upper‐crown leaves accords with better performance (with respect to dry pupal weight and relative growth rate) on the foliage; (iii) better performance is explained by a higher nitrogen content and N/C ratio of the upper‐crown foliage; and (iv) the effects of leaf quality on performance differ between sexes.     

不同生境栓皮栎天然更新幼苗植冠构型分析

栓皮栎存在于秦岭南坡的多种林分中,生活在不同生境中的个体往往形成不同的树冠形态和构型特征。为了说明不同生境条件下栓皮栎幼苗的植冠构型变化,采用典型抽样法,对秦岭南坡3种生境中(林冠下、林隙、林缘)的栓皮栎天然更新幼苗的侧枝、叶片特征及其空间分布进行了调查分析,结果表明:3种不同生境中栓皮栎幼苗植冠形态发生了明显的可塑性变化,(1)林冠下的幼苗明显为开阔型树冠,林隙和林缘处的幼苗树冠相对紧密;(2)幼苗的1级侧枝密度与分枝角度在3种生境下均差异显著(P0.05);从Ⅰ到Ⅳ层,林冠下幼苗的分枝角度在冠层内变化幅度不到5°,而林缘处幼苗的分枝角度变化高达40°;发生5个以上1级侧枝的概率以林冠下最大,为0.6;(3)从林缘、林隙到林冠下,幼苗的叶长、叶宽、单叶面积、叶面积指数逐渐降低,数量叶密度和比叶面积则逐渐增大,与其它两种生境相比,林冠下幼苗的叶片逐渐向树冠上层集中,且以更高序的侧枝为主要着生枝条;(4)林隙中栓皮栎幼苗的树高、地径明显优于林缘和林冠下,缩短了苗木进入主林层的时间,林隙对栓皮栎种群更新有利。在今后栓皮栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化的提供有利条件。     

Do Variations in Local Leaf Irradiance Explain Changes to Leaf Nitrogen within Row Maize Canopies?

Numerous studies have dealt with the relationship between leafnitrogen content and leaf irradiance. However, most of themrefer to dense stands presenting reduced horizontal heterogeneityof foliage distribution. Both gradients of leaf nitrogen andleaf irradiance related to canopy depth are significant undersuch conditions, and modelling radiative exchange using a turbid-mediumanalogy and dividing the canopy into vegetation layers is sufficient.Conversely, row crops such as maize are characterized by stronghorizontal heterogeneity of foliage distribution and the one-dimensional(1D) approach may be unsuitable. We thus modelled the three-dimensional(3D) geometry of maize canopies with varying densities and atdifferent developmental stages using plant digitizing underfield conditions. The nitrogen content per unit area of eachleaf part was obtained subsequently by nitrogen analysis. Wenext calculated radiative exchange using a 3D volume-based approachwithin the canopies in order to estimate local leaf irradianceon a daily integration scale. Vertical gradients in leaf nitrogencontent per unit area observed in dense stands during the vegetativephase corresponded largely to those reported in the literature.We also identified significant gradients in nitrogen contentalong the leaves, which had not before been clearly demonstrated.Our study shows that local light climate during plant developmentplays a major role in leaf nitrogen distribution and remobilization.Moreover, brutal plant thinning involves rapid changes in leafnitrogen partitioning. It is concluded that taking account ofthe 3D heterogeneity of nitrogen and irradiance distributionmay have implications for modelling crop photosynthesis andproduction. Copyright 1999 Annals of Botany Company 3D plant architecture, horizontal gradients in leaf nitrogen, leaf irradiance, leaf nitrogen content per unit area, maize, nitrogen partitioning, nitrogen remobilization, virtual plant, Zea mays L.     

Responses of sap flux and stomatal conductance of Pinus taeda L. trees to stepwise reductions in leaf area

Herbivory or artificial foliage removal has been shown to affect gas exchange and canopy water relations. In this study, canopy architecture and water relations in response to progressive defoliation were examined in a stand of 8-year-old loblolly pine (Pinus taeda L.) trees, a shade-intolerant, pioneer species common in the south-eastern USA. Sap flux was measured with constant heat sap flow gauges in order to estimate canopy stomatal conductance (Gs) while foliage in the 6 m high stand was harvested in 1 m increments from the bottom up. Leaf-level stomatal conductance and water potential data were also collected. Profiles of silhouette area ratio and specific leaf area showed no trends with crown height, reflecting an open canopy (leaf area index = 1.55). Therefore, short-term changes in Gs with foliage removal were attributed to hydraulic effects rather than influences of changes in mean microclimate conditions on Gs of remaining foliage. A large increase in Gs was observed during the 6 h pruning period which fully compensated for the reductions in foliage area down to 45%. Canopy stomatal conductance and whole plant liquid phase conductance as calculated from sap flux were both influenced by the rate of growth as indicated by the annual basal area increment.     

Growth analysis ofQuercus serrata seedlings withinMiscanthus sinensis grass canopies differing in light availability

The effects of different light regimes on the survival, growth and morphology ofQuercus serrata seedlings were studied in canopies ofMiscanthus sinensis. The seedlings of various ages (0–3 yr) were grown in three light regimes: under a denseM. sinensis canopy (TG plot) receiving 2.5%–8.7% of full sunlight, under a relatively sparse canopy (SG plot) receiving 3.8%–16.1% of light and in an adjacent open site (NG plot). There was a little difference in the survival ofQ. serrata seedlings among the three plots. Height and diameter of stem and total leaf area of the seedlings were significantly lower in the shadier plots. However, the first (bottom) flush of the stem was significantly longer in the TG plot than in the NG and SG plots. Total dry weights of individual 1- and 2-yr-oldQ. serrata seedlings in the TG plot were reduced to about one-twelfth of those in the NG plot. Although the relative proportion in dry weight of each organ did not differ significantly among the plots, leaf area ratio, specific leaf area and stem height per unit dry weight were significantly higher in shadier plots. The leaf area per unit stem height was increased considerably in the sunnier plots.     

辽东栎林内不同小生境下幼树植冠构型分析

以黄土高原黄龙山林区辽东栎林内3个小生境（林下、林隙、林缘）下辽东栎天然更新幼树为研究对象,采用典型抽样法对辽东栎幼树侧枝、叶片和树冠的空间分布状况以及生物量分配状况进行调查分析,探讨微生境与幼树植冠构型特征的关系,明确辽东栎幼树对不同小生境的适应策略,为栎林经营和林分结构优化提供理论依据。结果显示：（1）3种生境下辽东栎幼树构型发生了可塑性变化,林下幼树树冠层次比较单一,林隙与林缘的幼树树冠层次更加丰富。（2）由林下至林缘,幼树的树高、枝下高呈逐渐减小的趋势,而地径变化趋势与之相反;幼树的冠幅、树冠面积、树冠率呈先增加后减小的趋势,并且林下与林隙、林缘的差异显著;幼树的总体分枝率、逐步分枝率、枝径比呈先增加后减小的趋势。（3）3种生境下,幼树的一级枝的枝长、直径与倾角随着树高的增加而呈减小的趋势,但3种生境的差异不显著;林下一级枝主要分布在冠层中上部,而林隙与林缘一级枝主要分布在冠层中下、中上部。（4）由林下至林缘幼树叶长、叶宽、单叶面积和比叶面积逐渐降低,而单株叶数、叶总面积、叶面积指数呈先增大后减小趋势;与其他2种生境相比林下叶片分布趋于冠层上部。（5）幼树地上部分生物量中林下主干生物量占83%,枝和叶生物量只占17%;而林隙与林缘虽然各部位生物量有所差异但比例基本一致,其中主干占66%左右,枝和叶生物量占34%左右。研究表明,林隙生境下幼树的构型优于林缘和林下生境,在今后栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化提供有利条件。     

Canopy structure and leaf nitrogen distribution in a stand of Lysimachia vulgaris L. as influenced by stand density

Summary A hypothesis that a dense stand should develop a less uniform distribution of leaf nitrogen through the canopy than an open stand to increase total canopy photosynthesis was tested with experimentally established stands of Lysimachia vulgaris L. The effect of stand density on spatial variation of photon flux density, leaf nitrogen and specific leaf weight within the canopy was examined. Stand density had little effect on the value of the light extinction coefficient, but strongly affected the distribution of leaf nitrogen per unit area within a canopy. The open stand had more uniform distribution of leaf nitrogen than the dense stand. However, different light climates between stands explained only part of the variation of leaf nitrogen in the canopy. The specific leaf weight in the canopy increased with increasing relative photon flux density and with decreasing nitrogen concentration.     

Plastic leaf morphology in three species of Quercus: The more exposed leaves are smaller,more lobated and denser

Phenotypic plasticity and developmental instability in leaf traits are common in oak species but the role of environmental factors is not well understood. To decipher possible correlations between different leaf traits and effects of the position of leaves within the tree canopy, we quantified the plasticity of three leaf traits of 30 trees of Quercus alba L., Quercus palustris Muench and Quercus velutina Lam. We hypothesized that trees could modify the shape of their leaves for better adaptation to the variable microclimate within the canopy. Our results demonstrated that the south and north outer leaves were significantly smaller, more lobed and denser than those situated in the inner canopy. The order of leaves on the branch accounted for the plasticity of leaf traits in Q. alba only. Plasticity of lobing in Q. alba and Q. velutina depended on the height of the trees. We detected fluctuating asymmetry (FA) in all three species, but the source of variation depended on branch position in Q. velutina only. FA was more pronounced in north-facing leaves. Plasticity of the leaf traits ranged from small to medium. Plasticity of leaf area and leaf mass per area (LMA) depended on the branch position. However, the plasticity of lobation was not affected by the location of a branch within the tree canopy. Quercus alba and Q. palustris had similar plastic responses but the plasticity of Q. velutina was significantly smaller. We concluded that individual plants detect and cope with environmental stress through vegetative organ modification.     

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.     

Do the capacity and kinetics for modification of xanthophyll cycle pool size depend on growth irradiance in temperate trees?

The present study investigated the interaction of growth irradiance (Q_int) with leaf capacity for and kinetics of adjustment of the pool size of xanthophyll cycle carotenoids (sum of violaxanthin, antheraxanthin and zeaxanthin; VAZ) and photosynthetic electron transport rate (J_max) after changes in leaf light environment. Individual leaves of lower‐canopy/lower photosynthetic capacity species Tilia cordata Mill. and upper canopy/higher photosynthetic capacity species Populus tremula L. were either illuminated by additional light of 500–800 µmol m^?2 s^?1 for 12 h photoperiod or enclosed in shade bags. The extra irradiance increased the total amount of light intercepted by two‐fold for the upper and 10–15‐fold for the lower canopy leaves, whereas the shade bags transmitted 45% of incident irradiance. In control leaves, VAZ/area, VAZ/Chl and J_max were positively associated with leaf growth irradiance (Q_int). After 11 d extra illumination, VAZ/Chl increased in all cases due to a strong reduction in foliar chlorophyll, but VAZ/area increased in the upper canopy leaves of both species, and remained constant or decreased in the lower canopy leaves of T. cordata. The slope for VAZ/area changes with cumulative extra irradiance was positively associated with Q_int only in T. cordata, but not in P. tremula. Nevertheless, all leaves of P. tremula increased VAZ/area more than the most responsive leaves of T. cordata. Shading reduced VAZ content only in P. tremula, but not in T. cordata, again demonstrating that P. tremula is a more responsive species. Compatible with the hypothesis of the role of VAZ in photoprotection, the rates of photosynthetic electron transport declined less in P. tremula than in T. cordata after the extra irradiance treatment. However, foliar chlorophyll contents of the exposed leaves declined significantly more in the upper canopy of P. tremula, which is not consistent with the suggestion that the leaves with the highest VAZ content are more resistant to photoinhibition. This study demonstrates that previous leaf light environment may significantly affect the adaptation capacity of foliage to altered light environment, and also that species differences in photosynthetic capacity and acclimation potentials importantly alter this interaction.     

基于树形结构的木棉叶功能性状差异性研究

叶功能性状不仅反应植物对资源的利用能力,还涉及植物对自身结构和环境的生存适应策略。以不同生长阶段的木棉为研究对象,于2019年12月实地测量树高、冠幅等5个主要树形因子和采样测定叶面积、比叶面积、叶干物质含量、叶片含水量等14个叶功能性状,分析其在不同生长阶段的变化规律和相互关系,探讨叶功能性状对树形结构的响应。结果表明：不同生长阶段的木棉叶功能性状之间存在差异（P<0.05）,且表现出一定的相关性（P<0.05,P<0.01）;木棉叶片的面积、长度、宽度、周长、鲜重、饱和重、干重、比叶质量和干物质含量等功能性状是随生长阶段变化的主要指标。影响木棉叶片各性状的树形因子也不相同,其中叶面积主要受到冠幅的影响,叶片鲜重和饱和重主要受到枝下高的影响,叶片干重和相对含水量主要受到树高的影响,比叶面积、叶片含水量和干物质含量主要受到树高、尖削度的影响,比叶质量主要受到胸径、尖削度、树高的影响。随着木棉的生长,叶功能性状表现出受树形结构的影响而具有较强的表型可塑性,有利于其适应内外环境的变化。因此,叶功能性状之间的变化差异和相关组合,反映了不同生长阶段的木棉对树形结构的适应性调整和对资源的利用策略,在一定程度上为探索西双版纳热带雨林区的生态保护和可持续发展提供相关科学依据。     

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.     

Differences in sapling architecture betweenFagus crenata andFagus japonica

The ecological significance of architectural patterns for saplings ofFagus crenata andFagus japonica co-occurring in a secondary oak forest were evaluated by comparing the size and shape of leaves, trunks and crowns.Fagus japonica saplings were different fromF. crenata saplings in some architectural properties: (i) the leaf area and specific leaf area were larger; (ii) the ratio of sapling height to trunk length was lower, indicating greater leaning of the trunk; and (iii) the projection area of the crown was larger and the leaf area index lower indicating less mutual shading of leaves. These architectural features indicated thatF. japonica saplings were more shade tolerant thanF. crenata andF. crenata saplings were superior toF. japonica for growth in height and could, therefore, utilize sunlight in the upper layer. An erect trunk inF. crenata and a leaning trunk inF. japonica may be important characteristics associated with the regenerations patterns of each species; regeneration from seedlings under canopy gaps in the former and vegetative regeneration by sprouting in the latter.