Leaf functional traits are used in modeling forest canopy photosynthesis (Ac) due to strong correlations between photosynthetic capacity, leaf mass per area (LMA) and leaf nitrogen per area (Narea). Vertical distributions of these traits may change over time in temperate deciduous forests as a result of acclimation to light, which may result in seasonal changes in Ac To assess both spatial and temporal variations in key traits, we measured vertical profiles of Narea and LMA from leaf expansion through leaf senescence in a sugar maple (Acer saccharum Marshall) forest. To investigate mechanisms behind coordinated changes in leaf morphology and function, we also measured vertical variation in leaf carbon isotope composition (δ(13)C), predawn turgor pressure, leaf water potential and osmotic potential. Finally, we assessed potential biases in Ac estimations by parameterizing models with and without vertical and seasonal Narea variations following leaf expansion. Our data are consistent with the hypothesis that hydrostatic constraints on leaf morphology drive the vertical increase in LMA with height early in the growing season; however, LMA in the upper canopy continued to increase over time during light acclimation, indicating that light is primarily driving gradients in LMA later in the growing season. Models with no seasonal variation in Narea overestimated Ac by up to 11% early in the growing season, while models with no vertical variation in Narea overestimated Ac by up to 60% throughout the season. According to the multilayer model, the upper 25% of leaf area contributed to over 50% of Ac, but when gradients of intercellular CO2, as estimated from δ(13)C, were accounted for, the upper 25% of leaf area contributed to 26% of total Ac Our results suggest that ignoring vertical variation of key traits can lead to considerable overestimation of Ac.© The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.

Twig cross-sectional area and the surface area of leaves borne on it are expected to be isometrically correlated across species (Corner's rules). However, how stable this relationship remains in time is not known. We studied inter- and intraspecific twig leaf area-cross-sectional area (la-cs) and other scaling relationships, including the leaf-shoot mass (lm-sm) scaling relationship, across a complete growing season. We also examined the influence of plant height, deciduousness and the inclusion of reproductive buds on the stability of the scaling relationships, and we discuss results from a functional perspective.We collected weekly current-year twigs of six Patagonian woody species that differed in growth form and foliar habit. We also used prominent inflorescences from Embothrium coccineum (Proteaceae) to assess whether reproductive buds alter the la-cs isometric relationship. Mixed effects models were fitted to obtain parameter estimates and to test whether interaction terms were non-significant (invariant) for the scaling relationships.The slope of the la-cs scaling relationship remained invariant across the growing season. Two species showed contrasting and disproportional (allometric) la-cs scaling relationships (slope ≠ 1). Scaling relationships varied significantly across growth form and foliar habit. The lm-sm scaling relationship differed between reproductive- and vegetative-origin twigs in E. coccineum, which was explained by a significantly lower leaf mass per area in the former.Although phenology during the growing season appeared not to change leaf-shoot scaling relationships across species, we show that scaling relationships departed from the general trend of isometry as a result of within-species variation, growth form, foliar habit and the type of twig. The identification of these functional factors helps to understand variation in the general trend of Corner's rules.© The Author(s) 2020. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.

Silviculture focuses on establishing forest stand conditions that improve the stand increment. Knowledge about the efficiency of an individual tree is essential to be able to establish stand structures that increase tree resource use efficiency and stand level production. Efficiency is often expressed as stem growth per unit leaf area (leaf area efficiency), or per unit of light absorbed (light use efficiency). We tested the hypotheses that: (1) volume increment relates more closely with crown light absorption than leaf area, since one unit of leaf area can receive different amounts of light due to competition with neighboring trees and self-shading, (2) dominant trees use light more efficiently than suppressed trees and (3) thinning increases the efficiency of light use by residual trees, partially accounting for commonly observed increases in post-thinning growth. We investigated eight even-aged Norway spruce ( (L.) Karst.) stands at Bärnkopf, Austria, spanning three age classes (mature, immature and pole-stage) and two thinning regimes (thinned and unthinned). Individual leaf area was calculated with allometric equations and absorbed photosynthetically active radiation was estimated for each tree using the three-dimensional crown model Maestra. Absorbed photosynthetically active radiation was only a slightly better predictor of volume increment than leaf area. Light use efficiency increased with increasing tree size in all stands, supporting the second hypothesis. At a given tree size, trees from the unthinned plots were more efficient, however, due to generally larger tree sizes in the thinned stands, an average tree from the thinned treatment was superior (not congruent in all plots, thus only partly supporting the third hypothesis).

快捷、准确地测定植物叶片尺度上的叶面积(LA)和叶干质量(LDM)对于探讨植物性状对气候变化的响应机制至关重要,但适于测定区域尺度上不同阔叶植物单个叶片LA和LDM的经验模型尚未提出.本研究以中国东北4个分布区阔叶红松林内的白桦、紫椴、山杨、枫桦、水曲柳和裂叶榆6种阔叶树种为研究对象,分别测定其不同冠层高度(上层、中层和下层)叶片的叶长、叶宽、叶厚、LA及LDM.以叶长与叶宽之比(叶长宽比)的中位数为标准将6种阔叶树分为两组,检验每组树种不同冠层高度对构建预测LA和LDM的经验模型是否存在显著影响;构建适于预测区域尺度上不同冠层单个叶片LA和LDM的经验模型,验证其预测精度;并进一步评估构建的经验模型预测其他区域相同阔叶树种LA和LDM的适用性.结果表明: 整体上6种阔叶树单个叶片的LA随冠层高度的降低呈显著增大趋势,而部分树种的LDM呈下降趋势;冠层高度对构建预测LA和LDM的经验模型具有显著影响;构建的经验模型预测两组阔叶树种不同冠层单个叶片LA和LDM的平均精度分别为95%和83%,且基于这些模型预测其他区域相应树种LA和LDM的平均精度分别为94%和80%,表明本研究构建的经验模型在我国东北区域具有普遍适用性.

Knowledge of variations in morphophysiological leaf traits with forest height is essential for quantifying carbon and water fluxes from forest ecosystems. Here, we examined changes in leaf traits with forest height in diverse tree species and their role in environmental acclimation in a tropical rain forest in Borneo that does not experience dry spells. Height-related changes in leaf physiological and morphological traits [e.g., maximum photosynthetic rate (Amax), stomatal conductance (gs), dark respiration rate (Rd), carbon isotope ratio (δ(13)C), nitrogen (N) content, and leaf mass per area (LMA)] from understory to emergent trees were investigated in 104 species in 29 families. We found that many leaf area-based physiological traits (e.g., A(max-area), Rd, gs), N, δ(13)C, and LMA increased linearly with tree height, while leaf mass-based physiological traits (e.g., A(max-mass)) only increased slightly. These patterns differed from other biomes such as temperate and tropical dry forests, where trees usually show decreased photosynthetic capacity (e.g., A(max-area), A(max-mass)) with height. Increases in photosynthetic capacity, LMA, and δ(13)C are favored under bright and dry upper canopy conditions with higher photosynthetic productivity and drought tolerance, whereas lower R d and LMA may improve shade tolerance in lower canopy trees. Rapid recovery of leaf midday water potential to theoretical gravity potential during the night supports the idea that the majority of trees do not suffer from strong drought stress. Overall, leaf area-based photosynthetic traits were associated with tree height and the degree of leaf drought stress, even in diverse tropical rain forest trees.

Leaf initiation at the shoot apical meristem involves a balance between cell proliferation and commitment to make primordia. Several genes, such as CLAVATA1, CLAVATA3, WUSCHEL, KNOTTED1, and PHANTASTICA, play key roles in these processes. When expressed in the leaf primordium, however, these 'meristem' genes can profoundly affect leaf shape and size, possibly by regulating hormone gradients and transport. The KNOTTED1-like genes are involved in regulating changes in hormonal levels. Recent studies have elaborated on the role that hormones, such as auxin, play in releasing biophysical constraints on leaf initiation and growth. Final leaf form is elaborated by a coordination of these hormonally regulated processes, cell division and cellular differentiation.

Photosynthetic acclimation of deciduous broad-leaved tree species was studied along a vertical gradient within the canopy of a multi-species deciduous forest in northern Japan. We investigated variations in (1) local light regime and CO2 concentration ([CO2]), and (2) morphological (area, thickness and area per mass), biochemical (nitrogen and chlorophyll concentrations) and physiological (light-saturated photosynthetic rate) attributes of leaves of seven major species on three occasions (June, August and October). We studied early successional species, alder (Alnus hirsuta (Spach) Rupr.) and birch (Betula platyphylla var. japonica (Miq.) Hara); gap phase species, walnut (Juglans ailanthifolia Carrière) and ash (Fraxinus mandshurica var. japonica Rupr.); mid-successional species, basswood (Tilia japonica (Miq.) Simonk.) and elm (Ulmus davidiana var. japonica (Rehd.) Nakai); and the late-successional species, maple (Acer mono Bunge). All but maple initiated leaf unfolding from the lower part of the crown. The [CO2] within the vertical profile ranged from 320-350 ppm in the upper canopy to 405-560 ppm near the ground. The lowest and highest ambient [CO2] occurred during the day and during the night, respectively. This trend was observed consistently during the summer, but not when trees were leafless. Chlorophyll concentration was positively related to maximum photosynthetic rate within, but not among, species. Leaf senescence started from the inner part of the crown in alder and birch, but started either in the outer or top portion of the canopy of ash, basswood and maple. Chlorophyll (Chl) to nitrogen ratio in leaves increased with decreasing photon flux density. However, Chl b concentration in all species remained stable until the beginning of leaf senescence. Maximum photosynthetic rates observed in sun leaves of early successional species, gap phase or mid-successional species, and late successional species were 12.5-14.8 micromol m(-2) s(-1), 4.1-7.8 micromol m(-2) s(-1) and 3.1 micromol m(-2) s(-1), respectively.

比叶质量(LMA)是叶片经济型谱中最基础的叶功能性状, 也是重要的复合型结构参数。LMA不仅与植物的许多生理反应密切相关, 而且能定量测定光合产物在单位叶面积的投入, 因而被认为是反映植物生态策略的重要指标。目前, 有关LMA的深入研究已在植物生态学、农学、林学、植物生理学领域全面展开。该文系统阐述了LMA在叶片整体、组织、细胞三个水平的结构解析和计算方法; 重点分析了LMA对光合作用的影响; 讨论了LMA的内在遗传差异以及外部的环境胁迫因子(温度、水分、光照)对LMA的影响, 以期梳理比叶质量研究的思路、策略和方法, 为今后的研究提供借鉴和参考。

Leaf architecture, stand leaf area index and canopy light interception were studied in 13 poplar clones growing in a second rotation of a coppice plantation, to determine the role of leaf architectural attributes on canopy light-harvesting efficiency and to assess biomass investment in leaf support tissue. Stand leaf area index (L) varied from 2.89 to 6.99, but L was only weakly associated with canopy transmittance (TC). The weak relationship between TC and L was a result of a higher degree of foliage aggregation at larger values of L, leading to lower light-interception efficiency in stands with greater total leaf area. We observed a strong increase in leaf aggregation and a decrease in light-harvesting efficiency with decreasing mean leaf petiole length (PL) but not with leaf size, possibly because, in cordate or deltoid poplar leaves, most of the leaf area is located close to the petiole attachment to the lamina. Although PL was the key leaf characteristic of light-harvesting efficiency, clones with longer petioles had larger biomass investments in petioles, and there was a negative relationship between PL and L, demonstrating that enhanced light harvesting may lead to an overall decline in photosynthesizing leaf surface. Upper-canopy leaves were generally larger and had greater dry mass (MA) and nitrogen per unit area (NA) than lower-canopy leaves. Canopy plasticity in MA and NA was higher in clones with higher foliar biomass investment in midrib, and lower in clones with relatively longer petioles. These relationships suggest that there is a trade-off between photosynthetic plasticity and biomass investment in support, and also that high light-harvesting efficiency may be associated with lower photosynthetic plasticity. Our results demonstrate important clonal differences in leaf aggregation that are linked to leaf structure and biomass allocation patterns within the leaf.

To determine the role of leaf mechanical properties in altering foliar inclination angles, and the nutrient and carbon costs of specific foliar angle variation patterns along the canopy, leaf structural and biomechanical characteristics, biomass partitioning into support, and foliar nitrogen and carbon concentrations were studied in the temperate deciduous species Liriodendron tulipifera L., which possesses large leaves on long petioles. We used beam theory to model leaf lamina as a uniform load, and estimated both the lamina and petiole flexural stiffness, which characterizes the resistance to bending of foliar elements at a common load and length. Petiole and lamina vertical inclination angles with respect to horizontal increased with increasing average daily integrated photon flux density (Q ). Yet, the light effects on lamina inclination angle were primary determined by the petiole inclination angle. Although the petioles and laminas became longer, and the lamina loads increased with increasing Q, the flexural stiffness of both lamina and petiole increased to compensate for this, such that the lamina vertical displacement was only weakly related to Q. In addition, increases and decreases in the petiole inclination angle with respect to the horizontal effectively reduced the distance of lamina load from the axis of rotation, thereby reducing the bending moments and lamina inclination due to gravity. We demonstrate that large investments, up to 30% of total leaf biomass, in petiole and large veins are necessary to maintain the lamina at a specific position, but also that light has no direct effect on the fractional biomass investment in support. However, we provide evidence that apart from light availability, structural and chemical characteristics of the foliage may also be affected by water stress, magnitude of which scales positively with Q.

Extensive within-canopy light gradients importantly affect the photosynthetic productivity of leaves in different canopy positions and lead to light-dependent increases in foliage photosynthetic capacity per area (AA). However, the controls on AA variations by changes in underlying traits are poorly known. We constructed an unprecedented worldwide database including 831 within-canopy gradients with standardized light estimates for 304 species belonging to major vascular plant functional types, and analyzed within-canopy variations in 12 key foliage structural, chemical and physiological traits by quantitative separation of the contributions of different traits to photosynthetic acclimation. Although the light-dependent increase in AA is surprisingly similar in different plant functional types, they differ fundamentally in the share of the controls on AA by constituent traits. Species with high rates of canopy development and leaf turnover, exhibiting highly dynamic light environments, actively change AA by nitrogen reallocation among and partitioning within leaves. By contrast, species with slow leaf turnover exhibit a passive AA acclimation response, primarily determined by the acclimation of leaf structure to growth light. This review emphasizes that different combinations of traits are responsible for within-canopy photosynthetic acclimation in different plant functional types, and solves an old enigma of the role of mass- vs area-based traits in vegetation acclimation.© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.

Foliar biomass investment in support and assimilative compartments was studied in four temperate deciduous tree species along a natural light gradient across the canopy. The species ranked according to shade tolerance as Betula pendula Roth. < Populus tremula L. < Fraxinus excelsior L. < Tilia cordata Mill. Long-term light conditions at sampling locations were characterized as seasonal mean integrated quantum flux density (Q(int), mol m(-2) day(-1)) estimated by a method combining hemispherical photography and light measurements with quantum sensors. Leaf morphology was altered by Q(int) in all species. Both lamina and petiole dry mass per lamina area (LMA and PMA, respectively) increased with increasing Q(int). Shade-tolerant species had lower LMA at low Q(int) than shade-intolerant species; however, PMA was not related to shade tolerance. Across species, the ratio of petiole dry mass to lamina dry mass (PMR) varied from 0.07 to 0.21. It was independent of Q(int) in the simple-leaved species, but decreased with increasing Q(int) in the compound-leaved F. excelsior, which also had the largest foliar biomass investment in petioles. Differences in leaf mass and area, ranging over four orders of magnitude, provided an explanation for the interspecific variability in PMR. Species with large leaves also had greater biomass investments in foliar support than species with smaller leaves. This relationship was similar for both simple- and compound-leaved species. There was a negative relationship between PMR and petiole N concentration, suggesting that petioles had greater carbon assimilation rates and paid back a larger fraction of their construction cost in species with low PMR than in species with high PMR. This was probably the result of a negative relationship between PMR and petiole surface to volume ratio. Nevertheless, petioles had lower concentrations of mineral nutrients than laminas. Across species, the ratio of petiole N to lamina N varied from only 3 to 6%, demonstrating that petiole costs are less in terms of nutrients than in terms of total biomass, and that the petiole contribution to carbon assimilation is disproportionately lower than that of the lamina contribution.

The implications of extensive variation in leaf size for biomass distribution between physiological and support tissues and for overall leaf physiological activity are poorly understood. Here, we tested the hypotheses that increases in leaf size result in enhanced whole-plant support investments, especially in compound-leaved species, and that accumulation of support tissues reduces average leaf nitrogen (N) content per unit dry mass (N(M)), a proxy for photosynthetic capacity. Leaf biomass partitioning among the lamina, mid-rib and petiole, and whole-plant investments in leaf support (within-leaf and stem) were studied in 33 simple-leaved and 11 compound-leaved species. Support investments in mid-ribs and petioles increased with leaf size similarly in simple leaves and leaflets of compound leaves, but the overall support mass fraction within leaves was larger in compound-leaved species as a result of prominent rachises. Within-leaf and within-plant support mass investments were negatively correlated. Therefore, the total plant support fraction was independent of leaf size and lamina dissection. Because of the lower N(M) of support biomass, the difference in N(M) between the entire leaf and the photosynthetic lamina increased with leaf size. We conclude that whole-plant support costs are weakly size-dependent, but accumulation of support structures within the leaf decreases whole-leaf average N(M), potentially reducing the integrated photosynthetic activity of larger leaves.

More than 5,000 measurements from 1,943 plant species were used to explore the scaling relationships among the foliar surface area and the dry, water, and nitrogen/phosphorus mass of mature individual leaves. Although they differed statistically, the exponents for the relationships among these variables were numerically similar among six species groups (ferns, graminoids, forbs, shrubs, trees, and vines) and within 19 individual species. In general, at least one among the many scaling exponents was <1.0, such that increases in one or more features influencing foliar function (e.g., surface area or living leaf mass) failed to keep pace with increases in mature leaf size. Thus, a general set of scaling relationships exists that negatively affects increases in leaf size. We argue that this set reflects a fundamental property of all plants and helps to explain why annual growth fails to keep pace with increases in total body mass across species.

叶大小的变化是许多因素综合作用的结果, 对叶大小优化机制的研究有助于我们更好地理解植物的适应进化和生活史策略。该研究通过对浙江省清凉峰常绿阔叶混交林中的19个常绿阔叶物种和30个落叶阔叶物种叶水平上的相关性状进行分析, 探讨叶内生物量分配策略对叶大小优化的限制性影响。研究结果显示: 无论叶大小用面积还是质量表示, 常绿物种和落叶物种均呈现出叶内生物量分配到支撑结构的比例随着叶大小的增加而增加的规律, 这主要是由叶柄大小与叶片大小之间显著的异速生长关系导致的。这种异速生长关系在常绿物种和落叶物种中普遍存在。然而, 由于常绿物种对叶柄具有较高的机械以及抵抗冰冻栓塞等不利环境的需求, 在某一给定的叶面积下, 常绿物种比落叶物种具有更高的叶柄生物量投资。这些结果表明: 作为整个植株支撑投资的一个重要组成部分, 叶内支撑投资所占的生物量比例对叶大小的优化具有一定的限制性影响。

An assemblage of tree species with different crown properties creates heterogeneous environments at the canopy level. Changes of functional leaf traits are expected, especially those related to light interception and photosynthesis. Chlorophyll a fluorescence (ChlF) properties in dark-adapted leaves, specific leaf area, leaf nitrogen content (N) and carbon isotope composition (δ13C) were measured on Picea abies (L.) H.Karst., Pinus sylvestris L. and Betula pendula Roth. in monospecific and mixed boreal forests in Europe, in order to test whether they were affected by stand species richness and composition. Photosynthetic efficiency, assessed by induced emission of leaf ChlF, was positively influenced in B. pendula by species richness, whereas P. abies showed higher photosynthetic efficiency in monospecific stands. Pinus sylvestris had different responses when it coexisted with P. abies or B. pendula. The presence of B. pendula, but not of P. abies, in the forest had a positive effect on the efficiency of photosynthetic electron transport and N in P. sylvestris needles, and the photosynthetic responses were positively correlated with an increase of leaf δ13C. These effects on P. sylvestris may be related to high light availability at the canopy level due to the less dense canopy of B. pendula. The different light requirements of coexisting species was the most important factor affecting the distribution of foliage in the canopy, driving the physiological responses of the mixed species. Future research directions claim to enhance the informative potential of the methods to analyse the responses of pure and mixed forests to environmental factors, including a broader set of plant species' functional traits and physiological responses.© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.

不同物种间的功能性状差异是自然生态系统中物种共存的基础, 而物种内个体间的性状变异对物种的共存和分布同样具有重要作用。本文以湖北星斗山自然保护区亚热带常绿落叶阔叶混交林内28种主要树种(通过物种多度排序获得, 其中常绿和落叶树种各14种)为研究对象, 探讨不同叶习性树种的4种功能性状(比叶面积、叶干物质含量、叶面积和比茎密度)在种间和种内的差异程度。结果表明: (1)常绿和落叶树种在4种功能性状上均存在显著差异, 常绿树种的比叶面积和叶面积显著低于落叶树种, 但叶干物质含量和比茎密度则显著高于落叶树种; (2)比叶面积的变化主要来源于叶习性(57.49%), 叶面积变化主要来源于种间(66.80%)和种内变异(27.52%), 叶干物质含量的变化主要来源于种间(38.12%)和种内(33.88%)变异, 但比茎密度的变化主要来源于种内变异(51.50%), 其次为种间变异(32.52%); (3)常绿和落叶树种种间水平的性状相关性可能掩盖各功能性状之间的相关性。种内变异能够显著影响群落间的植物功能性状差异, 但不同功能性状的种内变异程度存在差异。

The majority of variation in six traits critical to the growth, survival and reproduction of plant species is thought to be organised along just two dimensions, corresponding to strategies of plant size and resource acquisition. However, it is unknown whether global plant trait relationships extend to climatic extremes, and if these interspecific relationships are confounded by trait variation within species. We test whether trait relationships extend to the cold extremes of life on Earth using the largest database of tundra plant traits yet compiled. We show that tundra plants demonstrate remarkably similar resource economic traits, but not size traits, compared to global distributions, and exhibit the same two dimensions of trait variation. Three quarters of trait variation occurs among species, mirroring global estimates of interspecific trait variation. Plant trait relationships are thus generalizable to the edge of global trait-space, informing prediction of plant community change in a warming world.

We explored the impact of canopy position on leaf respiration (R) and associated traits in tree and shrub species growing in a lowland tropical rainforest in Far North Queensland, Australia. The range of traits quantified included: leaf R in darkness (RD) and in the light (RL; estimated using the Kok method); the temperature (T)-sensitivity of RD; light-saturated photosynthesis (Asat); leaf dry mass per unit area (LMA); and concentrations of leaf nitrogen (N), phosphorus (P), soluble sugars and starch. We found that LMA, and area-based N, P, sugars and starch concentrations were all higher in sun-exposed/upper canopy leaves, compared with their shaded/lower canopy and deep-shade/understory counterparts; similarly, area-based rates of RD, RL and Asat (at 28 °C) were all higher in the upper canopy leaves, indicating higher metabolic capacity in the upper canopy. The extent to which light inhibited R did not differ significantly between upper and lower canopy leaves, with the overall average inhibition being 32% across both canopy levels. Log-log RD-Asat relationships differed between upper and lower canopy leaves, with upper canopy leaves exhibiting higher rates of RD for a given Asat (both on an area and mass basis), as well as higher mass-based rates of RD for a given [N] and [P]. Over the 25-45 °C range, the T-sensitivity of RD was similar in upper and lower canopy leaves, with both canopy positions exhibiting Q10 values near 2.0 (i.e., doubling for every 10 °C rise in T) and Tmax values near 60 °C (i.e., T where RD reached maximal values). Thus, while rates of RD at 28 °C decreased with increasing depth in the canopy, the T-dependence of RD remained constant; these findings have important implications for vegetation-climate models that seek to predict carbon fluxes between tropical lowland rainforests and the atmosphere. © The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.

叶片性状-环境关系对于预测气候变化对植物的影响至关重要。该研究以青藏高原东缘常见阔叶木本植物为研究对象, 从47个样点采集了332个物种共666个种群的叶片, 测量了15个叶片性状, 调查了该区域木本植物叶片性状的变异程度, 并从种内和种间水平探讨了叶片性状对环境的响应及适应策略。结果表明, 反眏叶片大小的性状均具有较高的变异, 其中, 叶片面积是变异程度最大的性状。除气孔密度外, 大多数叶片性状与海拔显著相关。气候是叶片性状变异的重要驱动因素, 3.3%-29.5%的叶片性状变异由气候因子组合解释。其中, 气温对叶片性状变异解释度最高, 日照时间能解释大部分叶片性状的变异, 而降水量对叶片性状变异的解释度相对较小。与环境(海拔和气候因子)显著相关的叶片性状在种内明显少于种间水平, 可能是植物性状之间的协同变化与权衡使种内性状变异比较小, 从而减弱了种内叶片性状与环境因子的相关性。研究结果总体表明,叶片性状与木本植物对环境的适应策略密切相关, 植物通过选择小而厚的叶片和较短的叶柄以适应高海拔的 环境。