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.     

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”     

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     

Testing the generality of the ‘leafing intensity premium’ hypothesis in temperate broad-leaved forests: a survey of variation in leaf size within and between habitats

Because leaf size scales negatively and isometrically with leaf number per shoot size (leafing intensity) in woody species, and because most tree and shrub species have small leaves, Kleiman and Aarssen (J Ecol 95:376–382, 2007) recently proposed that natural selection favors high leafing intensity resulting in small leaves, i.e., the leafing-intensity-premium hypothesis. However, empirical evidence for or against this hypothesis is still lacking. In addition, this hypothesis has not been examined in the context of how leaf size varies among habitats. To fill this void, we investigated leaf size frequency distributions of woody species from temperate China and explored the relationships among leaf mass, leaf number, and stem mass of current-year shoots of 133 woody species at low and high altitudes of three mountain ranges. The scaling relationships between leaf size and leafing intensity (leaf number per stem mass) were determined using both standardized major axis regression analyses and phylogenetically independent comparative techniques. In light of the leafing-intensity-premium hypothesis, we made three predictions: (1) leaf size frequency distributions should be right-skewed for each local study area and for the entire study region, (2) leafing intensities at different altitudes at different sites should differ while leafing intensities at comparable altitudes should be similar baring large taxonomic differences among sites, and (3) that leafing intensity should be higher for any given leaf size in habitats with small-leaved species. Significant negative and isometric scaling relationships between leaf size and leafing intensity were found to be consistently conserved independent of habitat type, both across species and across correlated evolutionary divergences. Within each mountain range or across the entire study region, leaf size frequency distributions were right-skewed, in accordance with our prediction. However, leafing intensity was smaller for any given leaf size at the altitude with smaller leafed species than for altitudes characterized by large leafed species, i.e., altitudes characterized by species with small leaves did not have consistently higher leafing intensities than other altitudes on each mountain range. Our analyses therefore indicate the direct adaptive value of leaf size but not the selective advantage in high leafing intensity as posited by the leafing-intensity-premium hypothesis. We suggest that this hypothesis explains less about the variation of leaf size among different habitats as it does about variation within habitats, i.e., the relative importance of the adaptive significance of leafing intensity and leaf size can and does vary with habitats.     

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)...     

我国温带山地森林48种常见树种叶片重量-出叶强度的关系

叶片是植物的主要光合器官, 其质量与数量的权衡关系体现植物对环境的适应策略。在全球气候变化的背景下, 研究叶片质量与数量关系有助于理解植物对环境变化的响应趋势。该研究应用标准化主轴回归方法, 探讨了我国温带山地森林中48个常见树种的单叶干重与出叶强度的权衡关系。结果表明, 所有物种以及落叶阔叶林、常绿和落叶阔叶树种、单叶以及亚冠层阔叶树种的单叶干重与出叶强度表现为异速生长关系; 针叶林、针阔混交林、常绿及落叶针叶树种、复叶以及冠层阔叶树种则表现为等速生长关系。研究结果表明, 叶大小和出叶强度并无恒定的权衡关系。     

THE COST OF MERISTEM LIMITATION IN POLYGONUM ARENASTRUM: NEGATIVE GENETIC CORRELATIONS BETWEEN FECUNDITY AND GROWTH

Growth and reproduction in higher plants depend on meristems, which have three developmental fates. A meristem can become reproductive, but doing so terminates its activity, it can differentiate vegetatively, or it can remain quiescent for extended periods. The first two fates are mutually exclusive, and only the second leads to the production of additional meristems for subsequent growth and reproduction. In Polygonum arenastrum (frequently referred to as P. aviculare in North American Floras), an annual species lacking quiescent meristems, a quantitative genetic analysis of inbred full-sibling families revealed genetic variation in the developmental pattern of axillary meristem commitment to vegetative growth versus reproduction. Developmental variation resulted in family differences in the age of first reproduction, in age-specific fecundity and growth, and in final plant size and reproductive output. Furthermore, there were strong negative genetic correlations between age-specific growth and fecundity. Early commitment of meristems to reproduction favors high early fecundity, but reduces the number of meristems available for vegetative differentiation, and leads to lowered growth rates and fecundity later in life, when meristems are limiting. Conversely, meristem commitment to vegetative growth early in life results in low early fecundity but high late fecundity and growth. Meristem limitation, like resource limitation, is a proximate mechanism that generates trade-offs between life history traits. Differences between meristem limitation and resource limitation are discussed. Meristem limitation leads automatically to a senescent life history because of the determinate fate of reproductive meristems. Developmental characters were also found to be genetically correlated with metamer characters (leaf size, internode length) and seed size in this selfing species. The pattern of correlation is suggestive of selection for particular suites of life history and morphological characters.     

Axillary meristem development in Arabidopsis thaliana

Axillary shoot apical meristems initiate post-embryonically in the axils of leaves. Their developmental fate is a main determinant of the final plant body plan. In Arabidopsis, usually a single axillary meristem initiates in the leaf axil even though there is developmental potential for formation of multiple branches. While the wild-type plants rarely form multiple branches in the leaf axil, tfl1-2 plants regularly develop two or more branches in the axils of the rosette leaves. Axillary meristem formation in Arabidopsis occurs in two waves: an acropetal wave forms during plant vegetative development, and a basipetal wave forms during plant reproductive development. We report here the morphological and anatomical changes, and the STM expression pattern associated with the formation of axillary and accessory meristems during Arabidopsis vegetative development.     

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...     

barren inflorescence2 regulates axillary meristem development in the maize inflorescence

Organogenesis in plants is controlled by meristems. Shoot apical meristems form at the apex of the plant and produce leaf primordia on their flanks. Axillary meristems, which form in the axils of leaf primordia, give rise to branches and flowers and therefore play a critical role in plant architecture and reproduction. To understand how axillary meristems are initiated and maintained, we characterized the barren inflorescence2 mutant, which affects axillary meristems in the maize inflorescence. Scanning electron microscopy, histology and RNA in situ hybridization using knotted1 as a marker for meristematic tissue show that barren inflorescence2 mutants make fewer branches owing to a defect in branch meristem initiation. The construction of the double mutant between barren inflorescence2 and tasselsheath reveals that the function of barren inflorescence2 is specific to the formation of branch meristems rather than bract leaf primordia. Normal maize inflorescences sequentially produce three types of axillary meristem: branch meristem, spikelet meristem and floral meristem. Introgression of the barren inflorescence2 mutant into genetic backgrounds in which the phenotype was weaker illustrates additional roles of barren inflorescence2 in these axillary meristems. Branch, spikelet and floral meristems that form in these lines are defective, resulting in the production of fewer floral structures. Because the defects involve the number of organs produced at each stage of development, we conclude that barren inflorescence2 is required for maintenance of all types of axillary meristem in the inflorescence. This defect allows us to infer the sequence of events that takes place during maize inflorescence development. Furthermore, the defect in branch meristem formation provides insight into the role of knotted1 and barren inflorescence2 in axillary meristem initiation.     

Developmental disaster1: A novel mutation causing defects during vegetative and inflorescence development in maize (Zea mays, Poaceae)

Axillary meristems, which give rise to branches and flowers, play a critical role in plant architecture and reproduction. To understand how axillary meristems initiate, we have screened for mutants with defects in axillary meristem initiation to uncover the genes controlling this process. These mutants, called the barren class of mutants in maize (Zea mays), have defects in axillary meristem initiation during both vegetative and reproductive development. Here, we identify and characterize a new member of the barren class of mutants named Developmental disaster1 (Dvd1), due to the pleiotropic effects of the mutation. Similar to the barren mutants, Dvd1 mutants have fewer branches, spikelets, florets, and floral organs in the inflorescence due to defects in the initiation of axillary meristems. Furthermore, double mutant analysis with teosinte branched1 shows that dvd1 also functions in axillary meristems during vegetative development. However, unlike the barren mutants, Dvd1 mutants are semidwarf due to the production of shorter internodes, and they produce leaves in the inflorescence due to the outgrowth of bract leaf primordia. The suite of defects seen in Dvd1 mutants, together with the genetic interaction of Dvd1 with barren inflorescence2, suggests that dvd1 is a novel regulator of axillary meristem and internode development.     

Meristem aging is not responsible for age-related changes in growth and abscisic acid levels in the Mediterranean shrub, Cistus clusii

To obtain new insights into the mechanisms underlying aging in perennials, we measured abscisic acid levels, growth and other stress indicators in leaves of Cistus clusii Dunal plants of different ages grown under Mediterranean field conditions. Recently emerged leaves from 9-year-old plants were compared to those of 1-year-old plants (obtained from cuttings from 9-year-old plants) to evaluate the effects of meristem aging on plant aging. Rooting and successful establishment of the cuttings allowed us to compare the physiology of plants with old meristems, but of different size. Plants obtained from cuttings were rejuvenated, with new leaves displaying a higher leaf area and chlorophyll content, but smaller leaf mass per unit area ratios and endogenous abscisic acid levels than those of 9-year-old plants. A comparative study in 1-, 4- and 9-year-old plants revealed that abscisic acid levels increase during the early stages of plant life (with increases of 90% between 1- and 4-year-old plants), but then remain constant at advanced developmental stages (between 4- and 9-year-old plants). Although leaf biomass was 53% smaller in 9-year-old compared to 4-year-old plants, the dry matter produced per shoot apical meristem was equivalent in both plant groups due to an increased number of leaves per shoot in the former. It is concluded that (i) C. clusii plants maintain the capacity to rejuvenate for several years; (ii) newly emerged leaves accumulate higher amounts of abscisic acid during early stages of plant life, but the levels of this phytohormone later remain constant; and (iii) although plant aging leads to the production of smaller leaves, the amount of biomass produced per shoot apical meristem remains constant at advanced developmental stages.     

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。     

Herbivory could unlock mutations sequestered in stratified shoot apices of genetic mosaics

Many higher plants have shoot apical meristems that possess discrete cell layers, only one of which normally gives rise to gametes following the transition from vegetative meristem to floral meristem. Consequently, when mutations occur in the meristems of sexually reproducing plants, they may or may not have an evolutionary impact, depending on the apical layer in which they reside. In order to determine whether developmentally sequestered mutations could be released by herbivory (i.e., meristem destruction), a characterized genetic mosaic was subjected to simulated herbivory. Many plants develop two shoot meristems in the leaf axils of some nodes, here referred to as the primary and secondary axillary meristems. Destruction of the terminal and primary axillary meristems led to the outgrowth of secondary axillary meristems. Seed derived from secondary axillary meristems was not always descended from the second apical cell layer of the terminal shoot meristem as is expected for terminal and primary shoot meristems. Vegetative and reproductive analysis indicated that secondary meristems did not maintain the same order of cell layers present in the terminal shoot meristem. In secondary meristems reproductively sequestered cell layers possessing mutant cells can be repositioned into gamete-forming cell layers, thereby adding mutant genes into the gene pool. Herbivores feeding on shoot tips may influence plant evolution by causing the outgrowth of secondary axillary meristems.     

Convergence in leaf phenology traits of two understory ferns in the northwestern Iberian Peninsula

Aims Plants control leaf phenology to maximize annual photosynthetic product. Although ferns play an important ecological role in many habitats, especially forests, their phenology traits have been poorly studied. Here, we investigate the leaf phenology of two ferns of the forest understorey and analyse the relationship between the timing of leaf emergence and spore dispersal and the effect of between-year climatic variation.Methods We compared the leafing and sporing phenologies of two ferns with very large (>2 m), overwintering leaves: Culcita macrocarpa and Woodwardia radicans. We regularly monitored individuals of six populations in the northwestern Iberian Peninsula during a 3-year study. We studied eight phenology variables: leafing start date, leafing end date, leaf expansion time, number of new leaves per individual, between-individual synchrony, within-individual synchrony, percentage of fertile leaves and spore release date. We also determined leaf mass per area (LMA) and gathered data on air temperature and humidity.Important findings Both C. macrocarpa and W. radicans produce few leaves (~2 leaves individual^-1 year^-1), which expand simultaneously for a very long period (from midwinter to early summer), are retained for more than 1 year (37 and 19 months, respectively) and have relatively high LMAs. Such traits, together with large leaf size, have also been found in seed plants from the forest understorey and represent adaptations to this light-limited environment. Spores of both study ferns are simultaneously released in late winter, with little between-year variation caused by differences in air humidity. This remarkable similarity between species suggests that the convergence in timing of leaf emergence favours the convergence in timing of spore dispersal.     

Determination for inflorescence development is a stable state,separable from determination for flower development in Pisum sativum L. buds

Terminal meristems of Pisum sativum (garden pea) transit from vegetative to inflorescence development, and begin producing floral axillary meristems. Determination for inflorescence development was assessed by culturing excised buds and meristems. The first node of floral initiation (NFI) for bud expiants developing in culture and for adventitious shoots forming on cultured meristems was compared with the NFI of intact control buds. When terminal buds having eight leaf primordia were excised from plants of different ages (i.e., number of unfolded leaves) and cultured on 6-benzylaminopurine and kinetin-supplemented medium, the NFI was a function of the age of the source plant. By age 3, all terminal buds were determined for inflorescence development. Determination occurred at least eight nodes before the first axillary flower was initiated. Thus, the axillary meristems contributing to the inflorescence had not formed at the time the bud was explanted. Similar results were obtained for cultured axillary buds. In addition, meristems excised without leaf primordia from axillary buds three nodes above the cotyledons of age-3 plants gave rise to adventitious buds with an NFI of 8.3 ±0.3 nodes. In contrast seed-derived plants had an NFI of 16.5 ±0.2. Thus cells within the meristem were determined for inflorescence development. These findings indicate that determination for inflorescence development in P. sativum is a stable developmental state, separable from determination for flower development, and occurring prior to initiation of the inflorescence at the level of meristems.     

Shoot-level biomass allocation is affected by shoot type in Fagus sylvatica

Aims The present study aims (i) to examine if recently reported interspecific shoot-level biomass allocational trade-offs, i.e. isometric trade-offs between leaf mass (LM) and stem mass (SM) and between leaf size and leaf number, hold intraspecifically and (ii) to explore whether those scaling relationships are independent of shoot type (i.e. long vs. short shoots).Methods In order to address our questions, we used Fagus sylvatica saplings growing under a broad light range that were sampled in the Western Carpathians Mountains (Slovakia).Important findings We found that: (i) intraspecific shoot-level biomass allocational trade-offs differ from those reported interspecifically and that (ii) long and short shoots differ in biomass allocation scaling coefficients. Allometric relationships with slopes statistically smaller than 1.0 or higher than^-1.0, were found between SM and LM and between mean leafing intensity and individual leaf mass, respectively, in long shoots. In contrast, isometric scaling was found in short shoots. This suggests that leaf mass in short shoots is unaffected by shoot stem mass, in contrast to long shoots. Short shoots also had a larger fraction of biomass allocated to leaves. Beech shoots, as has been observed in other shoot dimorphic species, are specialized, with short shoots specializing in carbon gain and long shoots in space acquisition. A greater shift in LM than in SM among species during speciation shifting from allometric intraspecific relationships to an isometric interspecific scaling relationship between those traits could explain the discrepancies between the outputs of the present intraspecific study and others similar studies. This study draws attention to the importance of considering shoot types in future studies dealing with allocation rules in species with dimorphic shoots.