The leaf size/number trade-off in herbaceous angiosperms

Aims In this study, we examined the extent to which between-species leaf size variation relates to variation in the intensity of leaf production in herbaceous angiosperms. Leaf size variation has been most commonly interpreted in terms of biomechanical constraints (e.g. affected by plant size limitations) or in terms of direct adaptation associated with leaf size effects in optimizing important physiological functions of individual leaves along environmental gradients (e.g. involving temperature and moisture). An additional interpretation is explored here, where adaptation may be more directly associated with the number of leaves produced and where relatively small leaf size then results as a trade-off of high 'leafing intensity'—i.e. number of leaves produced per unit plant body size.Methods The relationships between mean individual leaf mass, number of leaves and plant body size were examined for 127 species of herbaceous angiosperms collected from natural populations in southern Ontario, Canada.Important findings In all, 88% of the variation in mean individual leaf mass across species, spanning four orders of magnitude, is accounted for by a negative isometric (proportional) trade-off relationship with leafing intensity. These results parallel those reported in recent studies of woody species. Because each leaf is normally associated with an axillary bud or meristem, having a high leafing intensity is equivalent to having a greater number of meristems per unit body size—i.e. a larger 'bud bank'. According to the 'leafing intensity premium' hypothesis, because an axillary meristem represents the potential to produce either a new shoot or a reproductive structure, high leafing intensity should confer greater architectural and/or reproductive plasticity (with relatively small leaf size required as a trade-off). This greater plasticity, we suggest, should be especially important for smaller species since they are likely to suffer greater suppression of growth and reproduction from competition within multi-species vegetation. Accordingly, we tested and found support for the prediction that smaller species have not just smaller leaves generally but also higher leafing intensities, thus conferring larger bud banks, i.e. more meristems per unit plant body size.     

Leaf size versus leaf numbertrade-offs in dioecious angiosperms

Aims We explore the possible role of leaf size/number trade-offs for the interpretation of leaf size dimorphism in dioecious plant species.Methods Total above-ground biomass (both male and female) for three herbaceous dioecious species and individual shoots (from both male and female plants) for three woody dioecious species were sampled to record individual leaf dry mass, number of leaves, dry mass of residual above-ground tissue (all remaining non-leaf biomass), number of flowers/inflorescences (for herbaceous species) and number of branches.Important findings For two out of three woody species and two out of three herbaceous species examined, male plants produced smaller leaves but with higher leafing intensity—i.e. more leaves per unit of supporting (residual) shoot tissue or plant body mass—compared with females. Male and female plants, however, did not differ in shoot or plant body mass or branching intensity. We interpret these results as possible evidence for a dimorphic leaf deployment strategy that promotes both male and female function, respectively. In male plants, capacity as a pollen donor may be favored by selection for a broadly spaced floral display, hence favoring relatively high leafing intensity because this provides more numerous axillary meristems that can be deployed for flowering, thus requiring a relatively small leaf as a trade-off. In one herbaceous species, higher leafing intensity in males was associated with greater flower production than in females. In contrast, in female plants, selection favors a relatively large leaf, we propose, because this promotes greater capacity for localized photosynthate production, thus supporting the locally high energetic cost of axillary fruit and seed development, which in turn requires a relatively low leafing intensity as a trade-off.     

呼伦贝尔草原植物叶片大小与数量的权衡关系及其内在机制

叶片大小与数量的权衡关系是植物生长策略的基础,研究叶片大小与数量权衡关系的内在机制对于深刻理解植物生长策略具有重要意义.本研究以单个叶片干重表示叶大小,以单位茎上的叶片数量表示出叶强度,采用标准主轴回归分析方法研究呼伦贝尔草原植物叶片大小与数量的权衡关系以及内在机制.结果表明:陈旗嵯岗(典型草原)和陈旗八一(草甸草原)...     

Application of leaf size and leafing intensity scaling across subtropical trees

Understanding the scaling between leaf size and leafing intensity (leaf number per stem size) is crucial for comprehending theories about the leaf costs and benefits in the leaf size–twig size spectrum. However, the scaling scope of leaf size versus leafing intensity changes along the twig leaf size variation in different leaf habit species remains elusive. Here, we hypothesize that the numerical value of scaling exponent for leaf mass versus leafing intensity in twig is governed by the minimum leaf mass versus maximum leaf mass (M _min versus M _max) and constrained to be ≤−1.0. We tested this hypothesis by analyzing the twigs of 123 species datasets compiled in the subtropical mountain forest. The standardized major axis regression (SMA) analyses showed the M _min scaled as the 1.19 power of M _max and the ‐α (−1.19) were not statistically different from the exponents of M _min versus leafing intensity in whole data. Across leaf habit groups, the M _max scaled negatively and isometrically with respect to leafing intensity. The pooled data''s scaling exponents ranged from −1.14 to −0.96 for M _min and M _max versus the leafing intensity based on stem volume (LIV). In the case of M _min and M _max versus the leafing intensity based on stem mass (LIM), the scaling exponents ranged from −1.24 to −1.04. Our hypothesis successfully predicts that the scaling relationship between leaf mass and leafing intensity is constrained to be ≤−1.0. More importantly, the lower limit to scaling of leaf mass and leafing intensity may be closely correlated with M _min versus M _max. Besides, constrained by the maximum leaf mass expansion, the broad scope range between leaf size and number may be insensitive to leaf habit groups in subtropical mountain forest.     

No evidence of a generalized potential ‘cost’ of apical dominance for species that have strong apical dominance

尚无证据表明顶端优势强的物种存在广义顶端优势潜在“成本”     

Morphometric analysis of nymphalid butterfly wings: number,size and arrangement of scales,and their implications for tissue‐size determination

Animal body size and tissue size depend on genetic and environmental factors, but the precise mechanisms of how tissue size is determined in proportion to body size remain unknown. Here we focused on wings from three nymphalid butterflies, Junonia orithya (Linnaeus), Vanessa cardui (Linnaeus) and Danaus chrysippus (Linnaeus) (Lepidoptera: Nymphalidae), to examine the contributions of the number and size of scales to macroscopic structures, represented by wing compartments, and to investigate the positional dependence of scale size, density and arrangement. The whole wing area and wing compartment area exhibited a high correlation in all three species. Similarly, the wing compartment area and the blue or orange area showed a high correlation in three species, indicating isometric relationships among wings of different sizes. However, only in J. orithya, the blue area was highly correlated with the number of constituent scales and, to a lesser extent, with scale size. In contrast, reasonable correlations were obtained between the blue or orange area and the number of rows in all three species. These results suggest that variations of the background area accompany changes in the number of scales through changes in the number of rows. In a background region of the compartment, scale size gradually decreased and scale density increased from the proximal to the distal side in all three species. Our findings suggest that butterfly wing tissue size is determined primarily by the number of scale cells and secondarily by the size change of scale cells before or during the period of row arrangement.     

清凉峰不同海拔木本植物小枝内叶大小-数量权衡关系

权衡关系是生活史对策理论的基础, 叶大小-数量的权衡关系对理解叶大小进化具有重要的意义。该研究以单叶面积和单叶片干重表示叶大小, 用小枝干重和小枝茎干重表示小枝大小, 采用标准化主轴估计(standardized major axis estimation, SMA)和系统独立比较分析(phylogenetically independent contrast analysis, PIC)的方法, 对浙江省清凉峰自然保护区3个不同海拔落叶阔叶木本植物当年生小枝内的叶大小与数量间的关系进行研究。结果显示, 无论叶大小和小枝大小是用面积或干重表示, 在每个海拔, 叶大小与出叶强度均存在显著的等速负相关关系, 表明在落叶阔叶木本植物中发现的叶大小与出叶强度之间的权衡关系在不同生境物种中是普遍存在的, 植物在叶大小方面的种间变化, 可能不是自然选择的直接产物, 而是叶片数量变化权衡关系的一个副产物。不同海拔间的比较显示, 高海拔物种的叶面积或干重与出叶强度相关关系的y轴截距比中、低海拔物种小, 表明在出叶强度一定的情况下, 高海拔物种比低海拔物种具有更小的叶大小。与高海拔物种相比, 中海拔物种的共同斜率沿共同主轴有一个向上的位移, 表明中海拔物种比高海拔物种具有更大的叶大小, 但出叶强度更小。这些结果表明生境对叶大小-数量的权衡关系具有显著的影响, 中海拔生境具有更适宜植物生长的气候及养分条件, 而高海拔的低温等不利影响使得叶片变小。     

Ecological implications of a flower size/number trade-off in tropical forest trees

BACKGROUND: In angiosperms, flower size commonly scales negatively with number. The ecological consequences of this trade-off for tropical trees remain poorly resolved, despite their potential importance for tropical forest conservation. We investigated the flower size number trade-off and its implications for fecundity in a sample of tree species from the Dipterocarpaceae on Borneo. METHODOLOGY/PRINCIPAL FINDINGS: We combined experimental exclusion of pollinators in 11 species, with direct and indirect estimates of contemporary pollen dispersal in two study species and published estimates of pollen dispersal in a further three species to explore the relationship between flower size, pollinator size and mean pollen dispersal distance. Maximum flower production was two orders of magnitude greater in small-flowered than large-flowered species of Dipterocarpaceae. In contrast, fruit production was unrelated to flower size and did not differ significantly among species. Small-flowered species had both smaller-sized pollinators and lower mean pollination success than large-flowered species. Average pollen dispersal distances were lower and frequency of mating between related individuals was higher in a smaller-flowered species than a larger-flowered confamilial. Our synthesis of pollen dispersal estimates across five species of dipterocarp suggests that pollen dispersal scales positively with flower size. CONCLUSIONS AND THEIR SIGNIFICANCE: Trade-offs embedded in the relationship between flower size and pollination success contribute to a reduction in the variance of fecundity among species. It is therefore plausible that these processes could delay competitive exclusion and contribute to maintenance of species coexistence in this ecologically and economically important family of tropical trees. These results have practical implications for tree species conservation and restoration. Seed collection from small-flowered species may be especially vulnerable to cryptic genetic erosion. Our findings also highlight the potential for differential vulnerability of tropical tree species to the deleterious consequences of forest fragmentation.     

The generality of leaf size versus number trade-off in temperate woody species

Background and Aims

Trade-offs are fundamental to life-history theory, and the leaf size vs. number trade-off has recently been suggested to be of importance to our understanding leaf size evolution. The purpose of the present study was to test whether the isometric, negative relationship between leaf size and number found by Kleiman and Aarssen is conserved between plant functional types and between habitats.

Methods

Leaf mass, area and number, and stem mass and volume of current-year shoots were measured for 107 temperate broadleaved woody species at two altitudes on Gongga Mountain, south-west China. The scaling relationships of leaf size (leaf area and mass) vs. (mass- and volume-based) leafing intensity were analysed in relation to leaf habit, leaf form and habitat type. Trait relationships were determined with both a standardized major axis method and a phylogenetically independent comparative method.

Key Results

Significant negative, isometric scaling relationships between leaf size and leafing intensity were found to be consistently conserved across species independent of leaf habit, leaf form and habitat type. In particular, about 99 % of the variation in leaf mass across species could be accounted for by proportional variation in mass-based leafing intensity. The negative correlations between leaf size and leafing intensity were also observed across correlated evolutionary divergences. However, evergreen species had a lower y-intercept in the scaling relationships of leaf area vs. leafing intensity than deciduous species. This indicated that leaf area was smaller in the evergreen species at a given leafing intensity than in the deciduous species. The compound-leaved deciduous species were observed usually to have significant upper shifts along the common slopes relative to the simple-leaved species, which suggested that the compound-leaved species were larger in leaf size but smaller in leafing intensity than their simple counterparts. No significant difference was found in the scaling relationships between altitudes.

Conclusions

The negative, isometric scaling relationship between leaf size and number is largely conserved in plants, while the leaf size vs. number trade-off can be mediated by leaf properties. The isometry of the leaf size vs. number relationship may simply result from a biomass allocation trade-off, although a twig size constraint may provide an alternative mechanism.Key words: Allometry, trade-off, leafing intensity, leaf size, leaf habit, leaf form     

武夷山不同海拔黄山松枝叶大小关系

对武夷山自然保护区不同海拔黄山松叶片面积、叶片数量、小枝长度及小枝直径等性状进行测定,分析不同海拔黄山松枝-叶大小间的权衡关系.结果表明: 随海拔升高,黄山松小枝的叶片数量、小枝长度、小枝直径、出叶强度及茎截面积逐渐增大,单叶面积呈逐渐减小趋势;不同海拔黄山松小枝出叶强度与单叶面积均呈显著负相关,不同海拔黄山松小枝茎截面积与总叶面积呈显著正相关;不同海拔黄山松小枝长度、小枝直径与出叶强度呈显著负相关,与单叶面积、叶片数量及总叶面积均呈显著正相关.为提高竞争优势或是资源利用效率,低海拔黄山松倾向于在短枝上着生量少但单叶面积大的针叶,而高海拔黄山松趋向于在长枝上着生量大但单叶面积小的针叶,这体现出不同海拔梯度黄山松小枝的资源利用策略及枝叶间生物量分配的权衡机制.     

不同海拔典型竹种枝叶大小异速生长关系

为了分析竹子枝-叶大小间的权衡关系,本研究对武夷山不同海拔典型竹种(毛竹、箬竹、肿节少穗竹、毛竿玉山竹和武夷山玉山竹)小枝的叶片总质量、茎质量、单叶质量和出叶强度等性状进行测定.结果表明: 随海拔升高,5个竹种间小枝上总叶质量与茎质量的异速生长指数呈显著下降趋势.竹种内,毛竹、箬竹和肿节少穗竹总叶质量与茎质量在不同海拔上均拥有共同异速生长指数(分别为0.94、0.85、0.84).毛竿玉山竹和武夷山玉山竹的叶茎质量也存在共同异速生长指数(0.79).除武夷山玉山竹外,竹子单叶质量与出叶强度之间均呈显著的负相关关系.5个竹种的单叶质量和出叶强度之间存在共同异速生长指数-1.12.总之,竹类植物的小枝总体上倾向于在低海拔环境中着生更多的叶片,而在高海拔生境下则投资更多的生物量到茎的构造上.尽管竹种间小枝的茎投资随海拔升高而增加,但其基于茎质量的出叶强度策略取决于叶片大小的构建而不是海拔生境差异.     

长期增温对树线交错带岷江冷杉幼苗异龄叶大小与出叶强度关系的影响

全球气候变暖强烈影响树线交错带植物的生活史策略,异龄叶大小-出叶强度权衡关系是常绿植物生活史策略的重要内容。以川西树线交错带的岷江冷杉（Abies faxoniana）幼苗为例,研究气候变暖对异龄叶大小与出叶强度关系的影响。通过开顶箱（Open-top chamber, OTC）对川西王朗自然保护区树线交错带的岷江冷杉进行模拟增温,采用标准化主轴估计（Standardized major axis estimation, SMA）方法研究了叶大小（单叶质量、单叶面积）与出叶强度（基于茎生物量、茎体积）间异速生长关系对长期增温的响应及其年际变化。结果表明：使用不同参数表征叶大小与出叶强度得到的结果存在差异;多年生小枝上存在单叶质量-出叶强度的负等速权衡关系,共同主轴随小枝年龄增加而向下漂移;长期增温并不影响单叶质量与出叶强度的异速生长关系,不同年龄小枝的异速生长常数对增温具有差异性响应。增温处理中当年生小枝在相同单叶质量下的出叶强度更低,以换取叶片总数的增加,使小枝具有更大的可塑性而适应增温。本研究提供了岷江冷杉幼苗协调异龄叶大小与出叶强度从而适应长期增温的证据,为评估树木生长随气候变化而加速提供了理论参考。     

胡杨叶片大小与叶数量的权衡关系对地下水埋深的响应

本文以胡杨(Populus euphratica Oliv.)为研究对象,采用标准化主轴回归估计(SMA)方法探讨了种群叶片大小(单叶面积、单叶质量)与出叶强度(基于小枝质量、茎质量、茎体积)的异速生长关系对地下水埋深梯度(Groundwater depth,GWD)的响应及其变化规律.结果显示:随GWD增加,胡杨当年...     

Reproductive allometry in Malaysian rain forest trees: Biomechanics versus optimal allocation

Summary Quantitative predictions of reproductive allometry in iteroparous plants may be derived from two bodies of theory: biomechanics and optimal allocation theory. Biomechanical theory predicts allometric scaling exponents between reproductive (R) and vegetative (V) biomass in the range of 0.44–1.33, while very general models of life history evolution predictR–V exponents > 1 in all cases. These predictions are examined in light of allometric patterns of flower and fruit production in 32 species of Malaysian rain forest trees. Among these species the mean estimatedR–V exponents are in the range 1.8–2.0 for staminate flower, pistillate flower and fruit production. This range of exponent values provides unambiguous support for some of the general predictions of optimal allocation models, but not for biomechanical theory. Optimal allocation models also predict a positive relationship between species size andR–V slope and a positive relationship between species size andR–V intercept parameters. The latter, but not the former prediction is supported by the data.R–V allometries in sexes of dioecious species were also found to differ in intercept, though not slope, reflecting smaller sizes at reproductive onset in staminate trees. Further critical examinations of reproductive allometry are encouraged as a relatively unexplored avenue for increasing the contact of theory and data in studies of life history evolution in long-lived organisms such as tropical trees.     

Influence of leaf size on tree architecture: first branch height and crown dimensions in tropical rain forest trees

 First branch height is an important attribute of sapling architecture, as it defines the height at which prolonged lateral growth is possible. First branch height, measured on saplings of 70 species in tropical rain forests of Australia, Costa Rica, Panama, and Sabah, Malaysia, was highly correlated with leaf blade and petiole length. The observed relationship, first branch height ∝ blade length × (petiole length)^0.5, implies that the ratio of first branch height to blade length increases somewhat with increasing leaf size, among species with a given ratio of petiole to blade length. Orthotropic species, with more or less radially symmetric arrangements of leaves on ascending axes, had a mean first branch height of 7x that observed for plagiotropic species, with planar leaf arrangements. The greater first branch height of orthotropic species was associated with their larger leaves and longer petioles. Plagiotropic species had wider crowns than orthotropic species in the sapling stage, as assessed at the Costa Rican site. Thus, leaf dimensions influence the dynamics of crown construction (or visa versa), as well as affecting leaf energy balance and gas exchange. Received: 5 September 1997 / Accepted: 3 March 1998     

The trade-off between number and size of offspring in humans and other primates

Life-history theory posits a fundamental trade-off between number and size of offspring that structures the variability in parental investment across and within species. We investigate this 'quantity-quality' trade-off across primates and present evidence that a similar trade-off is also found across natural-fertility human societies. Restating the classic Smith-Fretwell model in terms of allometric scaling of resource supply and offspring investment predicts an inverse scaling relation between birth rate and offspring size and a (-1/4) power scaling between birth rate and body size. We show that these theoretically predicted relationships, in particular the inverse scaling between number and size of offspring, tend to hold across increasingly finer scales of analyses (i.e. from mammals to primates to apes to humans). The advantage of this approach is that the quantity-quality trade-off in humans is placed into a general framework of parental investment that follows directly from first principles of energetic allocation.     

东灵山不同林型五角枫叶性状异速生长关系随发育阶段的变化

分析植物叶片性状种内水平的异速生长关系有助于加深理解生长发育过程中叶片的资源利用模式.分析了东灵山3个主要林型(白桦林、胡桃楸林、辽东栎林)中五角枫成树、幼树、幼苗的叶面积、叶体积、叶含水量与叶干重之间的异速生长关系.结果表明:成树叶面积增长速度小于叶干重的增长速度,幼树、幼苗叶面积与叶干重保持同速增长;成树、幼树叶干重与叶体积保持同速增长,幼苗叶体积的增长速度超过了干重的增长速度;成树叶含水量的增长速度小于叶干重的增长速度,幼树、幼苗两性状间保持等速增长.叶含水量与叶干重的异速生长指数在不同的林型间有显著差异,白桦林叶含水量的增长速度小于叶干重的增长速度,其余两个林型均为等速增长趋势.这些结果揭示了不同发育阶段五角枫资源利用方式的转变.随叶干重的增加,成树将更多的叶生物量投资于同化和支持结构;幼树则保持对光合面积和光合同化结构的稳步投资;而幼苗主要投资于叶面积的增大.叶含水量与叶干重的异速曲线在不同林型间的差异说明叶片代谢活性相较于其他叶性状可塑性更高.     

The scaling of leaf area and mass: the cost of light interception increases with leaf size

For leaves, the light-capturing surface area per unit dry mass investment (specific leaf area, SLA) is a key trait from physiological, ecological and biophysical perspectives. To address whether SLA declines with leaf size, as hypothesized due to increasing costs of support in larger leaves, we compiled data on intraspecific variation in leaf dry mass (LM) and leaf surface area (LA) for 6334 leaves of 157 species. We used the power function LM=alpha LAbeta to test whether, within each species, large leaves deploy less surface area per unit dry mass than small leaves. Comparing scaling exponents (beta) showed that more species had a statistically significant decrease in SLA as leaf size increased (61) than the opposite (7) and the average beta was significantly greater than 1 (betamean=1.10, 95% CI 1.08-1.13). However, scaling exponents varied markedly from the few species that decreased to the many that increased SLA disproportionately fast as leaf size increased. This variation was unrelated to growth form, ecosystem of origin or climate. The average within-species tendency found here (allometric decrease of SLA with leaf size, averaging 13%) is in accord with concurrent findings on global-scale trends among species, although the substantial scatter around the central tendency suggests that the leaf size dependency does not obligately shape SLA. Nonetheless, the generally greater mass per unit leaf area of larger than smaller leaves directly translates into a greater cost to build and maintain a unit of leaf area, which, all else being equal, should constrain the maximum leaf size displayed.