古田山自然保护区四个常绿阔叶树种叶面积的估算模型

通过对浙江古田山自然保护区4个主要阔叶树种——甜槠、木荷、青冈和石栎的调查采样,测定了4个物种共1160张叶片的叶长、叶宽和叶面积,基于此构建4个物种叶面积与叶长和(或)叶宽的8种常用回归模型。结果表明,模型SA=a(LW)+b具有最高的决定系数(R~2=0.97-0.99)和最低的均方差(MSE=0.52-1.59);幂指数模型SA=aW~b是所有单变量模型中拟合效果最佳的模型。     

Characterization of a type II chlorophyll a/b-binding protein gene (Lhcb2*Pp1) in peach. II. mRNA abundance in developing leaves exposed to sun or shade

Leaf development of shoots exposed to full sunlight and shoots shaded by the canopy was followed in field-grown, mature peach trees (Prunus persica (L.) Batsch, cv. Loring) during the first half of the 1995 growing season. The architecture and size of shaded shoots and sun-exposed shoots differed significantly. Total number of leaves produced on shaded shoots was significantly less than on sun-exposed shoots throughout the season, and differences in leaf number between light conditions increased as the season progressed. The overall patterns of leaf development along sun-exposed and shaded shoots were qualitatively similar. The expression pattern of the type II chlorophyll a/b-binding protein gene, Lhcb2*Pp1, determined by RNA abundance in leaves at different positions along the shoot, was also similar between the two light conditions. The major difference between sun-exposed and shaded leaves was a lower abundance of Lhcb2*Pp1 RNA in mature, shaded leaves compared with sun-exposed leaves. Although the number of fruit per shoot was significantly lower on shaded shoots than on sun-exposed shoots, the rate of fruit drop was not substantially different during the growing season, indicating that quantitative differences in leaf initiation and growth caused by differences in light exposure did not adversely affect fruit retention. However, based on comparison with a previous study of leaf development in non-fruiting trees, reproductive development slowed the rate of vegetative growth without affecting the overall pattern of leaf development along the shoots.     

Allometric models for leaf area and leaf mass predictions across different growing seasons of elm tree (<Emphasis Type="Italic">Ulmus japonica</Emphasis>)

Convenient and effective methods to determine seasonal changes in individual leaf area (LA) and leaf mass (LM) of plants are useful in research on plant physiology and forest ecology. However, practical methods for estimating LA and LM of elm (Ulmus japonica) leaves in different periods have rarely been reported. We collected sample elm leaves in June, July and September. Then, we developed allometric models relating LA, LM and leaf parameters, such as leaf length (L) and width (W) or the product of L and W (LW). Our objective was to find optimal allometric models for conveniently and effectively estimating LA and LM of elm leaves in different periods. LA and LM were significantly correlated with leaf parameters (P < 0.05), and allometric models with LW as an independent variable were best for estimating LA and LM in each period. A linear model was separately developed to predict LA of elm leaves in June, July and September, and it yielded high accuracies of 93, 96 and 96%, respectively. Similarly, a specific allometric model for predicting LM was developed separately in three periods, and the optimal model form in both June and July was a power model, but the linear model was optimal for September. The accuracies of the allometric models in predicting LM were 88, 83 and 84% for June, July and September, respectively. The error caused by ignoring seasonal variation of allometric models in predicting LA and LM in the three periods were 1–4 and 16–59%, respectively.     

Leaf senescence and late-season net photosynthesis of sun and shade leaves of overstory sweetgum (Liquidambar styraciflua) grown in elevated and ambient carbon dioxide concentrations

We examined the effects of elevated CO2 concentration ([CO2]) on leaf demography, late-season photosynthesis and leaf N resorption of overstory sweetgum (Liquidambar styraciflua L.) trees in the Duke Forest Free Air CO2 Enrichment (FACE) experiment. Sun and shade leaves were subdivided into early leaves (formed in the overwintering bud) and late leaves (formed during the growing season). Overall, we found that leaf-level net photosynthetic rates were enhanced by atmospheric CO2 enrichment throughout the season until early November; however, sun leaves showed a greater response to atmospheric CO2 enrichment than shade leaves. Elevated [CO2] did not affect leaf longevity, emergence date or abscission date of sun leaves or shade leaves. Leaf number and leaf area per shoot were unaffected by CO2 treatment. A simple shoot photosynthesis model indicated that elevated [CO2] stimulated photosynthesis by 60% in sun shoots, but by only 3% in shade shoots. Whole-shoot photosynthetic rate was more than 12 times greater in sun shoots than in shade shoots. In senescent leaves, elevated [CO2] did not affect residual leaf nitrogen, and nitrogen resorption was largely unaffected by atmospheric CO2 enrichment, except for a small decrease in shade leaves. Overall, elevated [CO2] had little effect on the number of leaves per shoot at any time during the season and, therefore, did not change seasonal carbon gain by extending or shortening the growing season. Stimulation of carbon gain by atmospheric CO2 enrichment in sweetgum trees growing in the Duke Forest FACE experiment was the result of a strong stimulation of photosynthesis throughout the growing season.     

Effects of current-year and previous-year PPFDs on shoot gross morphology and leaf properties in Fagus japonica

We investigated how shoot gross morphology and leaf properties are determined in Fagus japonica Maxim., a deciduous species with flush-type shoot phenology, in which all leaves are produced in a single flush at the start of each season. We examined relationships between current-year shoot properties and local light environment in a 14-m tall beech tree growing in a deciduous forest. Leaf number (LN), total leaf area (TLA), and total leaf length (SL) of the current-year shoot increased with increasing photosynthetic photon flux density (PPFD). Leaf thickness, dry mass per leaf area and nitrogen content on a leaf area basis increased, whereas the chlorophyll/N ratio decreased with increasing PPFD. To separate the effects of current-year PPFD from those of previous year(s), we artificially shaded a part of the uppermost leaf tier. Reciprocal transfers of beech seedlings between controlled PPFD regimes were also made. Characteristics of shoot gross morphology such as LN, TLA and SL were largely determined by the PPFD of the previous year. The exception was the length of the longest "long shoots" with many leaves, in which elongation appeared to be influenced by both previous-year and current-year PPFD. In contrast, leaf properties were determined by current-year PPFD. The ecological implications of our findings are discussed.     

Gas exchange characteristics of mangosteen (Garcinia mangostana L.) leaves

We used the plastochron index to study the relationship between plant age, leaf age and development, and net photosynthesis of black cherry (Prunus serotina Ehrh.) seedlings. Leaf area and net photosynthesis were measured on all leaves >/= 75 mm of plants ranging in age from 7 to 20 plastochrons. Effects of plant developmental stage on leaf area and net photosynthesis were evaluated for leaves of differing age (horizontal series), leaves on plants of constant age (vertical series), and leaves of constant age (oblique series). Regression techniques were used to estimate leaf area from leaf blade dimensions. The best equations for predicting leaf area had R(2) values of 0.991-0.992 and used linear or logarithmic functions of both leaf length and width. Suitable, but less precise, equations with R(2) values of 0.946-0.962 were developed from either leaf length or leaf width. Leaf area development in black cherry seedlings was similar to that in other indeterminate species. Leaves of young plants reached full expansion at a lower leaf plastochron age than leaves of older plants. Maximum net photosynthesis per unit leaf area occurred 2-3 plastochrons before full leaf expansion. There was strong ontogenetic drift in net photosynthesis with leaf age; net photosynthesis decreased as plant age increased in leaves of the same plastochron age. Plots of the oblique series were particularly useful in providing information about interaction effects.     

Phenology and photosynthetic traits of short shoots and long shoots in Betula grossa

Leaf phenology, growth irradiance (i.e., photosynthetic photon flux (PPF) at the leaf surface) and photosynthetic capacity (A(area); measured at a PPF of 1000 micro mol m(-2) s(-1) and expressed on a leaf area basis) were investigated in early leaves (ELs) and late leaves (LLs) of Betula grossa Siebold & Zucc. trees. Early leaves first appeared on morphologically distinct long shoots and short shoots. The appearance of ELs, which was restricted to the bud break period, was followed by the successive appearance of LLs on long shoots only. Late leaves appeared successively until the middle of the growing season. Late leaves started to abscise around the middle of the growing season, whereas ELs on both long and short shoots did not abscise until near the end of the growing season. Solar irradiance was higher at the surface of LLs of late appearance than at the surface of either LLs of early appearance or ELs. Solar irradiance at the surface of ELs decreased after LLs appeared. In both long and short shoots, A(area) of ELs increased and then remained stable for 65-80 days before starting to decrease. Although A(area) was higher in LLs than in ELs for a short time in August, it started to decrease earlier in LLs than in ELs. Area-based nitrogen concentration (N(area)) was higher in LLs than in ELs after August. Although N(area) decreased slowly in ELs after August, it did not decrease in LLs. In both ELs and LLs, A(area)/N(area) decreased with time. The crown was thus characterized by a rapidly growing surface with young LLs having high A(area) and by shaded inner parts with ELs having stable low A(area).     

白刺叶片形态变化对叶片δ13C的影响

选取磴口、民勤唐古特白刺,敦煌东湖附近洪积扇中部、底部生长的泡泡刺为研究对象,通过叶片扫描与Image-Pro Plus6.0图像处理软件分析获得叶片形态参数(长、宽、周长和面积),应用同位素分析获得叶片δ13C值。结果表明:随着水分有效性的增加,白刺叶片面积具有增加趋势,叶片面积在增加过程中,叶片长、宽增幅具有非同步性;相同叶片宽度,生长于水分条件较好地区的唐古特白刺和泡泡刺叶片更长。虽然磴口、民勤及敦煌东湖附近洪积扇中部、底部水分环境存在较大的差异,但生长于这些区域的唐古特白刺及泡泡刺叶片δ13C值的差异不显著(P>0.05)。叶片周长/面积比是连接叶片形态变化与植物水分生理的重要因子;白刺叶面积增加过程中,叶片长度的优先增加有利于减缓周长/面积比的下降速率,从而使白刺保持较高的水分利用效率。     

Variation in Leaf Shapes of Nitraria Species and Effect on Leaf δ13C

Leaf shapes are not only the useful indicators in plant taxonomy,but also the important factors affecting energy and material exchange in leaves.In this paper,we collected and scanned the leaves of Nitraria tangutorum in Dengkou of Inner Mongolia Autonomous Region（the mean annual precipitation 145 mm） and Minqin of Gansu Province （the mean annual precipitation 115 mm） and N. sphaerocarpa in Dunhuang,and then analyzed leaf shape parameters with Image - Pro Plus6.0 image processing software and leafδ¹³C values in the isotope laboratory of the Chinese Academy of Forestry.The result showed that:1) as leaf area increased with increasing water availability the increases in the leaf length and width were asynchronously;2) with the same leaf width,the 1 eaves of N.tangutorum and N.sphaerocarpa were significantly longer in high water available conditions;and 3) although there were significantly differences in water availability between Dengkou and Minqin,as well as between the bottom and middle of the alluvial fan near the East Lake in Dunhuang,the leafδ¹³C values of N.tangutorum or N.sphaerocarpa were similar in different water conditions（P>0.05）.Our results suggested that the ratio of leaf perimeter to area would be an important factor which linked leaf shape to plant water physiology.During growing procedure of leaf area,leaf length increase was prior to its width to alleviate the reduction in ratio of perimeter to area and maintain water use efficiency of the plant.     

观赏植物叶面积测定及相关分析

采用扫描仪采集鹅掌柴、仙客来、三角梅、非洲茉莉、桂花和绿萝6种植物叶片图像,利用像素得到叶片真实面积,与叶面积仪测定的叶面积相比较,结果表明叶面积仪能够完全反映叶片的真实面积;确定了6种植物的叶面积、比叶面积、叶形指数和叶面积指数,并对叶面积与叶宽进行了相关性分析,结果表明叶面积和叶宽的相关性显著,进一步回归分析得出叶宽和叶面积的一元二次方程,即只需测量叶宽就可推算叶面积的简便方法;通过对比叶面积进行聚类分析,得出6种植物适宜的栽培环境,为合理管理观赏植物提供了可靠的参考依据。     

日本核用银杏枝叶生长及黄酮变化规律的研究

为 给银杏引 种和产 业化生产 提 供科 学依 据， 采用 日 本的 藤九 郎、 金兵 卫、 黄金 丸 和岭南４ 个大果 银杏的 标准 枝 和标 准 叶， 对 其 枝 叶生 产 及 黄硐 变 化规 律 进 行了 研 究。 结 果证 明，新梢长、 叶数 枝、叶柄 长、叶长 、叶 宽、 单叶 面 积、 单叶 干 重和 鲜 重 的年 变 化呈 Ｓ 型曲 线；新梢长 和叶数 枝分别 在７ 月４ 日和６ 月１４ 日达最 大值， 而 干重在７ 月底 达最 大值。 芽内 黄酮含量＞ 刚刚展开 的嫩叶 ＞ 成熟叶 ＞ 黄叶 ； ６ 月 初叶 内黄 酮含 量 下降 到原 初 的４０９ ％ ， 到１１ 月１６ 日又回升 到３０４ ％ ； 叶内黄酮 含量与 叶位及 叶龄 有关； 新梢 上部 叶及 幼 龄叶 单叶 面 积、干重、鲜 重低于下 部叶及老 龄叶， 但 黄酮含 量恰好相 反。     

Practical methods for non-destructive measurement of tree leaf area

In an agroforestry context, the knowledge of leaf area is an important parameter to take into consideration because tree foliage shades the intercrop. Single leaf measurement (for example leaf length and width) is a widely used method to estimate leaf area in a rapid non-destructive way. In this study, the objectives were to estimate the leaf area of different leaf sizes and shapes for Acacia mangium Willd. (Mimosaceae), Tectona grandis L. (Verbenaceae), Hevea brasiliensis Muell.-Arg. (Euphorbiaceae), and Swietenia macrophylla King. (Meliaceae) Indonesian agroforestry trees. In order to characterise leaf area in field conditions a digital photograph method was used. Leaf length was measured in order to build relationships between leaf length versus actual leaf area. Additional measurements obtained from the digital pictures such as lamina width, number of leaflets, leaflet length and width were recorded in order to test the benefit regarding leaf area estimation based only on the leaf length model. The combination of these different linear leaf measurements led to various degrees of precision of leaf area estimation. The different models can then be chosen according to a compromise between the accuracy of leaf area estimation and the time allocated to field measurements.     

Leaf hydraulic conductance in relation to anatomical and functional traits during Populus tremula leaf ontogeny

Leaf hydraulic conductance (K(leaf)) and several characteristics of hydraulic architecture and physiology were measured during the first 10 weeks of leaf ontogeny in Populus tremula L. saplings growing under control, mild water deficit or elevated temperature conditions. During the initial 3 weeks of leaf ontogeny, most measured characteristics rapidly increased. Thereafter, a gradual decrease in K(leaf) was correlated with a decrease in leaf osmotic potential under all conditions, and with increases in leaf dry mass per area and bulk modulus of elasticity under mild water deficit and control conditions. From about Week 3 onward, K(leaf) was 33% lower in trees subjected to mild water deficit and 33% higher in trees held at an elevated temperature relative to control trees. Mild water deficit and elevated temperature treatment had significant and opposite effects on most of the other characteristics measured. The ontogenetic maximum in K(leaf) was correlated positively with the width of xylem conduits in the midrib, but negatively with the overall width of the midrib xylem, number of lateral ribs, leaf dry mass per area and bulk modulus of elasticity. The ontogenetic maximum in K(leaf) was also correlated positively with the proportion of intercellular spaces and leaf osmotic potential, but negatively with leaf thickness, volume of mesophyll cells and epidermis and number of cells per total mesophyll cell volume, the closest relationships being between leaf osmotic potential and number of cells per total mesophyll cell volume. It was concluded that differences in protoplast traits are more important than differences in xylem or parenchymal cell wall traits in determining the variability in K(leaf) among leaves growing under different environmental conditions.     

Leaf exchange in a Mediterranean shrub: water, nutrient, non-structural carbohydrate and osmolyte dynamics

Leaf exchange is an abrupt phenological event that drastically modifies the morphology and physiology of the aerial portion of the plant. We examined if water and osmolyte differences between old leaves and new organs trigger leaf exchange, and whether the differences are closely linked to the resource resorption process in senescing leaves. We monitored concentrations of osmolyte, water, non-structural carbohydrate, nitrogen and potassium in senescing leaves and in emerging new leaves and inflorescences of a Mediterranean leaf exchanger (Cistus laurifolius L.) growing in NE Spain. Old leaves rehydrated markedly during most of the senescence process, which co-occurred with the extension of new shoots, suggesting the lack of a clear-cut switch in water supply from old to new organs. The accumulation of osmolytes in the early stage of leaf senescence might account for this rehydration. Osmolyte dynamics in old leaves depended largely on the progression of resource resorption from senescing organs but were mostly unrelated to water content during late senescence. We conclude that dehydration of old leaves is not a prerequisite for the triggering of leaf exchange. The finding that most nutrients and carbohydrates accumulated in new organs before senescing leaves massively exported resources, and the absence of relevant differences between the dynamics of old leaves at the base of inflorescences and those at the base of vegetative shoots, indicate that the nutrient and carbohydrate demands of new organs do not trigger leaf exchange.     

Structural variation in current-year shoots of broad-leaved evergreen tree saplings under forest canopies in warm temperate Japan

Stem length and leaf area of current-year shoots were measured in saplings of eight broad-leaved evergreen tree species growing under a forest canopy. Stem length varied over a range of one to two orders of magnitude within each species. In all species, both the number of leaves and the mean stem length between successive leaves were greater in longer shoots. Mean leaf size and stem length were not correlated in six of eight species, and only weakly positively correlated in the other two species. Thus, total leaf area per stem increased with stem length, but not in direct proportion: leaf area per stem length was smaller in shoots with long stems and larger in shoots with short stems. I conclude that the within-species variation in the leaf-stem balance of current-year shoots is related to variation in shoot functional roles, as has been observed for long and short shoots in many deciduous tree species: shoots with long stems are extension oriented and contribute to the framework of the crown, whereas shoots with short stems serve mainly for leaf display. Among species, large differences were found in the leaf area per stem length ratio. In the species with larger leaf area per stem length ratios, leaves had narrower blades or longer petioles, or both, resulting in a reduction of mutual shading among the leaves on the shoot.     

Interspecific variation of photosynthesis and leaf characteristics in canopy trees of five species of Dipterocarpaceae in a tropical rain forest

Photosynthetic rate, nitrogen concentration and morphological properties of canopy leaves were studied in 18 trees, comprising five dipterocarp species, in a tropical rain forest in Sarawak, Malaysia. Photosynthetic rate at light saturation (Pmax) differed significantly across species, varying from 7 to 18 micro mol m(-2) s(-1). Leaf nitrogen concentration and morphological properties, such as leaf blade and palisade layer thickness, leaf mass per area (LMA) and surface area of mesophyll cells per unit leaf area (Ames/A), also varied significantly across species. Among the relationships with leaf characteristics, Pmax had the strongest correlation with leaf mesophyll parameters, such as palisade cell layer thickness (r2 = 0.76, P < 0.001) and Ames/A (r2 = 0.73, P < 0.001). Leaf nitrogen concentration and Pmax per unit area also had a significant but weaker correlation (r2 = 0.46, P < 0.01), whereas Pmax had no correlation, or only weakly significant correlations, with leaf blade thickness and LMA. Shorea beccariana Burck, which had the highest P(max) of the species studied, also had the thickest palisade layer, with up to five or more layers. We conclude that interspecific variation in photosynthetic capacity in tropical rain forest canopies is influenced more by leaf mesophyll structure than by leaf thickness, LMA or leaf nitrogen concentration.     

Relationships between light, leaf nitrogen and nitrogen remobilization in the crowns of mature evergreen Quercus glauca trees

We estimated the amount of nitrogen (N) remobilized from 1-year-old leaves at various positions in the crowns of mature Quercus glauca Thunb. ex Murray trees and related this to the production of new shoots. Leaf N concentration on an area basis (Na) and total N (Nt= Na x lamina area of all leaves on a shoot) were related to photosynthetic photon flux (PPF) on the leaves of current-year and 1-year-old shoots. When new shoots (S02 shoots; flushed in 2002) flushed, only a portion of the leaves on the previous year's shoots (S01 shoots; flushed in 2001) were shed. After the S02 shoots flushed, S01 shoots were defined as 1-year-old shoots (S01* shoots). Both Na and Nt were positively correlated with PPF for S01 shoots, but not for S01* shoots. The fraction of remobilized N (% of the maximum Na in S01 leaves) from remaining leaves was 5-35%, with the fraction size being positively correlated with the number of S02 shoots on an S01* shoot (new shoot number). However, the mean fraction of remobilized N from fallen leaves was 45% and was unrelated to new shoot number. The total amount of N remobilized from both fallen and remaining leaves was 1-20 mg per S01* shoot. Total remobilized N was positively correlated with new shoot number. There was a statistically significant positive relationship between the light-saturated net photosynthetic rate on a leaf area basis (Amax) and Na for both S01* and S02 leaves. However, when we compared leaves with similar Na, Amax of S01* leaves was only half that of S02 leaves, indicating that 1-year-old leaves had lower instantaneous N-use efficiency (Amax per unit Na) than current-year leaves. Ratios of chlorophyll a:b and Rubisco:chlorophyll were lower in S01* leaves than in S02 leaves, indicating that 1-year-old leaves were acclimatized to lower light environments. Thus, in Q. glauca, the N allocation theory (i.e., that N is distributed according to local PPF) applied only to the current-year shoots. Although the amount of foliar N in 1-year-old shoots was not strongly affected by the PPF on 1-year-old leaves, it was affected by interactions with current-year shoots.     

Influence of shoot architectural position on shoot growth and branching patterns in Cleyera japonica

The influence of shoot architectural position on shoot growth and branching patterns was examined in saplings of Cleyera japonica Thunb. (Theaceae), an understory, broad-leaf evergreen woody species. Shoot length varied with branching order and the vertical position of the branch in the crown. In the upper crown, shoot length decreased with increasing branching order, whereas in the lower crown, differences in shoot length among branching orders were not significant. These results demonstrate that it is important to consider not only individual shoots, but also the relationships between shoots in terms of their architectural positions when studying the development of crown architecture in trees. Shoot branching patterns also varied with branching order and the vertical position of the branch in the crown. In the upper crown, branching was mainly sylleptic. In the middle of the crown, mainly proleptic branches were produced. In the lower crown, there was little branching. The importance of these trends in shoot growth and shoot branching patterns in terms of carbon production efficiency is discussed.     

霍城县大西沟野杏枝叶形态多样性

为给进一步开展野杏种质资源多样性及亲缘关系研究提供参考,选取新疆霍城大西沟不同区域分布的野杏单株,利用DPS7.05数据处理软件对枝叶形态性状指标进行聚类分析和主成分分析,探讨新疆野杏株间枝叶形态特征差异。聚类分析结果显示,系统聚类将62个野杏单株的15个枝叶形态性状分为4大类;主成分分析结果表明,前7个主成分的累计贡献率约为89.04%,影响较大的因子是叶片长度、叶片宽度、长宽比、叶柄长、叶面积、叶背绒毛、叶缘形态、新梢长度和粗度等。数量分类结果能反映出各单株间的遗传差异,大西沟野杏具有丰富的多样性,通过主成分分析选出了野杏分类的主要性状。通过聚类分析能清楚反映出所选野杏单株间的亲缘关系。     

Petiole length and biomass investment in support modify light interception efficiency in dense poplar plantations

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.