Adaxial–abaxial polarity: The developmental basis of leaf shape diversity

Leaves of flowering plants are diverse in shape. Part of this morphological diversity can be attributed to differences in spatiotemporal regulation of polarity in the upper (adaxial) and lower (abaxial) sides of developing leaves. In a leaf primordium, antagonistic interactions between polarity determinants specify the adaxial and abaxial domains in a mutually exclusive manner. The patterning of those domains is critical for leaf morphogenesis. In this review, we first summarize the gene networks regulating adaxial–abaxial polarity in conventional bifacial leaves and then discuss how patterning is modified in different leaf type categories. genesis 52:1–18, 2014. © 2013 The Authors. Genesis Published byWiley Periodicals, Inc.     

Determinants of some leaf characteristics of Australian mangroves

Leaf characteristics reflecting the size, lifespan (longevity), moisture content (degree of succulence) and complexity of structure of 20 mangrove species were studied over several years at 13 locations along the tropical and subtropical Australian coast. These characteristics were found to fall generally within the ranges of those for woody species from other ecosystems. With the exception of one species, it was found that leaf longevity was related inversely to leaf moisture content, increasing from nearly 6 months in more succulent species to over 2 years in less succulent species. This suggested that more succulent leaves are less complex in their structure because they have less well‐developed ability to compartmentalize salt. There was a tendency also for leaf longevity to increase in species with larger leaves. These findings were consistent with the general view for land plants that leaf longevity is greater in species that have developed tolerance to environmental stress, salt particularly in the case of mangroves. Leaf tissue in such species is more robust or complex and requires greater metabolic resources in its construction; the plant is then advantaged by retaining the tissue for longer periods. Classification of the species considered here, based on their leaf longevity, moisture content and complexity, identified phylogenetically related species groupings that reflected these leaf longevity effects.     

Evolution of leaf developmental mechanisms

Leaves are determinate organs produced by the shoot apical meristem. Land plants demonstrate a large range of variation in leaf form. Here we discuss evolution of leaf form in the context of our current understanding of leaf development, as this has emerged from molecular genetic studies in model organisms. We also discuss specific examples where parallel studies of development in different species have helped understanding how diversification of leaf form may occur in nature.     

Relationships between sclerophylly,leaf biomechanical properties and leaf anatomy in some Australian heath and forest species

ABSTRACT

A previous study of 19 south-east Australian heath and forest species with a range of leaf textures showed that they varied considerably in leaf biomechanical properties. By using an index of sclerophylly derived from botanists' rankings (botanists' sclerophylly index, BSI) we determined that leaves considered by botanists to be sclerophyllous generally had both high strength and work to fracture (particularly in punching and tearing tests), both at the level of leaf and per unit leaf thickness. In the current study we have shown that leaves from the same species also varied considerably in leaf specific mass (46–251 g m^-2), neutral detergent fibre concentration (20–59% on a dry weight basis) and in leaf anatomy. Multiple regression indicated a very strong correlation between BSI and the first two components of a principal components analysis (PCA) of leaf anatomy (R ² = 0.91). In addition, there was strong correlation between the first component of a PCA of the mechanical properties (correlated with BSI) and the two axes derived from anatomical characteristics (R ² = 0.66). The anatomical properties contributing most to the significant component axes were thickness of palisade mesophyll and upper cuticle (axis 1) and percentage fibre (neutral detergent fibre) and lower epidermis thickness (axis 2). However, whether these relationships are causal, or reflect correlations with characteristics not measured in this study, such as vascularization and sclerification, is not clear. At a finer scale, however, there is evidence that there are various ways to be sclerophyllous, both in terms of anatomical and mechanical properties. This is illustrated by comparison of two of the sclerophyllous species, Eucalyptus baxteri and Banksia marginata.     

Diurnal cycles of leaf extension in unsalinized and salinized Beta vulgaris

Abstract. Continuous high resolution measurement of sugar beet leaf extension over 5 d in growth chambers showed average leaf extension rates (LER) in darkness to be from three to six times those in light for plants growing in non-salinized media. The changes in LER in light-dark transitions occurred within seconds, a response which was more rapid than stomatal opening or closing. When the growth medium was salinized to 100 mol m⁻³ NaCl, LER's were reduced by about 50% in darkness and 90% in light, markedly increasing the ratio of dark to light LER.
A 2-d episode of root-zone salinity imposed midway through a 5-d period of measurement decreased LER and produced higher leaf temperatures. LER and diurnal leaf temperature patterns reverted to their pre-salinized levels when root-zone salinity was removed. Thus, the effects of short episodes of high sodium chloride in the growth medium appear to be reversible, suggesting a water stress mechanism of growth reduction rather than toxicity effects of salt.     

叶脉结构与功能及其对叶片经济谱的影响

叶脉由贯穿于叶肉内部的维管组织及其外围机械组织构成, 多样化的脉序及网络结构使叶脉系统发生变异和功能分化。该文综述了叶脉系统结构与功能的最新研究进展。通过聚焦叶脉分级系统的结构与功能及其在叶片经济谱(LES)中的重要性, 解释叶脉性状与其它叶片功能性状之间的关系及机制。不同等级叶脉在机械支撑与水分运输方面存在功能分化, 其中1-3级粗脉在维持叶片形状和叶表面积以及物理支撑方面发挥重要作用, 有利于维持叶片最大受光面积; 4级及以上细脉具有水分调节功能, 它们与气孔相互协调, 影响叶片水分运输、蒸腾散热和光合作用速率。叶片生长过程与叶脉发育的动态变化模式决定叶脉密度, 并影响叶脉密度与叶片大小之间的关系: 叶面积与粗脉密度呈显著负相关, 与粗脉直径呈显著正相关, 而与细脉密度无关。与叶脉性状相关的叶片经济谱框架模型预测, 叶脉密度较高的叶片寿命短、比叶重较小, 叶片最大碳同化速率、代谢速率以及资源获取策略潜力较高。     

Leaf traits indicate survival strategies among 42 dominant plant species in a dry, sandy habitat, China

The objective of this paper was to assess the congruency of leaf traits and soil characteristics and to analyze the survival strategies of different plant functional types in response to drought and nutrient-poor environ-ments in the southeastern Ke'erqin Sandy Lands in China. Six leaf traits-leaf thickness (TH), density (DN), specific leaf area (SLA), leaf dry weight to fresh weight ratio (DW/ FW), leaf N concentration (Nmass), and N resorption efficiency (NREmass)-of 42 plant species were investi-gated at four sites. The correlations between leaf traits and soil characteristics-organic C (OC), total N (TN), total P (TP), and soil moisture (SM)-were examined. We found that the six leaf traits across all the 42 species showed large variations and that DW/FW was negatively correlated with OC, TN, TP, and SM (P<0.05), while other leaf traits showed no significant correlations with soil characteristics. To find the dissimilarity to accommodate environment, a hierarchical agglomerative clustering analysis was made of all the species. All the species clustered into three groups except the Scutellaria baicalensis. Species of group Ⅲ might be most tolerant of an arid environment, and species of group Ⅱ might avoid nutrient stress in the nutrient-poor environment, while group Ⅰ was somewhat intermediate. Therefore, species from the different groups may be selected for use in vegetation restoration of different sites based on soil moisture and nutrient conditions.     

Leaf area estimation in a sugar beet cultivar by linear models

An indirect method of leaf area measurement for Rizor sugar beet cultivar was tested. Leaves were sampled during two growing seasons in a Randomised Complete Block Design experiment. For 2002 samplings, leaf area [cm²] was linearly correlated with maximum leaf width [cm] using all leaf samples (r ² = 0.83, p < 0.001) or using the means of the 8 sampling occasions (r ² = 0.97, p < 0.001). Correlations between leaf area and leaf mid vein length [cm] were weaker (r ² = 0.75, p < 0.001 and r ² = 0.93, p < 0. 001, respectively). For 2003 samplings, the area estimated by the equations was highly correlated to the measured leaf area.     

Hydraulic limitations imposed by crown placement determine final size and shape of Quercus rubra L. leaves

The canopies of large broad‐leaf trees exhibit significant heterogeneity in both micro‐environmental conditions and leaf morphology. Whether the visible differences in the size and shape of leaves from the top and bottom of the crown are determined prior to bud break or result from different patterns of leaf expansion is not known. Analysis of ontogenetic changes of both the degree of lobing and vein density in Quercus rubra demonstrates that leaves throughout the crown are identical in size and shape at the time of bud break. Morphological adaptation to the local micro‐environment takes place during the expansion phase and starts after the determination of the vascular architecture has been completed. Leaves from the bottom of the crown undergo greater expansion in the tissue close to the main veins than occurs either in the more peripheral tissue of the same leaf or anywhere in leaves from the top of the crown. This results in a water transport system that is well suited to the low evaporative rates near the bottom of the crown, but inadequate for the conditions found at the top of the tree. Acclimation of leaf form and function based upon differential expansion may be entirely driven by the local hydraulic demand during the expansion phase, resulting in leaf size and vein density being determined during development by the same hydraulic properties which will constrain the size of leaf that can be functionally supported at maturity.     

浙江天童常绿植物当年生与往年生叶片性状的变异与关联

植物叶片性状随叶龄的变化是植物生活史策略的体现, 反映了植物叶片的物质投资和分配方式。该研究通过在个体和物种2个水平, 比较浙江天童1 hm ²样地内常绿阔叶树种的平均叶面积(MLA)、比叶面积(SLA)和叶片干物质含量(LDMC)在当年生和往年生叶片间的差异和关联, 探究叶片物质分配策略在异龄叶间的变化, 并分析叶龄对植物叶片性状, 特别是叶片面积建成消耗的影响。结果显示: 1)在个体和物种水平上, MLA变异系数最大(个体: 79.5%; 物种: 66.5%), SLA次之(个体: 28.1%; 物种: 24.7%), LDMC较低(个体: 17.0%, 物种: 14.1%); 当年生叶片MLA、LDMC和SLA的变异系数均高于往年生叶片; 2)往年生叶MLA显著大于当年生叶(t = -38.53, p < 0.001), 往年生叶SLA显著小于当年生叶(t = 45.30, p < 0.001), 往年生叶LDMC显著大于当年生叶(t = -9.71, p < 0.001); 3)在个体水平, 当年生叶片MLA、SLA和LDMC值分别解释了往年生叶片MLA、SLA和LDMC变异的86%、48%和41%; 在物种水平, 当年生叶片MLA、SLA和LDMC值分别解释了往年生叶片MLA、SLA和LDMC变异的97%、83%和85%; 4) SLA在异龄叶间的变化表明, 与往年生叶片相比, 投资相同干物质, 当年生叶片可形成较大的叶面积, 其叶片面积建成消耗较小。研究认为, 植物叶性状在异龄叶间具有较大的变异性和关联性, 叶面积形成过程中生物量建成与消耗的协调可能影响植物叶片的发育。     

着生位置对水曲柳小叶性状变异及性状间相关性的影响

复叶植物相比单叶植物更具生长优势, 但复叶内部小叶性状及其相关关系是否受到着生位置影响尚未可知。该研究以东北典型复叶植物水曲柳(Fraxinus mandshurica)为研究对象, 测定复叶内部不同着生位置小叶的叶厚(LT)、叶面积(LA)、叶干物质含量(LDMC)、比叶面积(SLA)、叶氮含量(LNC)和叶磷含量(LPC), 分析上述6种小叶性状及其生长关系在复叶内部的变异, 并分别通过最小显著性差异(LSD)法以及标准化主轴(SMA)法检验着生位置对小叶性状及性状间生长关系是否存在显著影响。结果表明: (1) LT、LA、LDMC和LNC随小叶着生位置级别增加(从复叶顶端至复叶基部)呈减小趋势, 但SLA和LPC呈增大趋势。(2)复叶内部, LNC与SLA间以及LT与LDMC间表现为同速生长关系, LT、SLA、LPC 3个性状与LA间, SLA、LNC、LPC 3个性状与LDMC间以及LPC与LT间均表现为异速生长关系。(3)小叶着生位置对LA与LT、SLA、LPC之间的相关关系存在显著影响, LT、SLA与LA的斜率在三级小叶(复叶中部)附近达到最大值, LT、LPC与LA的斜率绝对值在六级小叶(复叶基部)处达到最小值。整体而言, 复叶内部小叶性状随着生位置存在一定变异规律, 小叶性状间多表现为异速生长关系, 且小叶性状间的生长关系一定程度上受着生位置的调控。     

Leaf demography of <Emphasis Type="Italic">Festuca pallens</Emphasis> in dry grassland communities

Leaf blade parameters and leaf demography of Festuca pallens Host were studied in two types of dry grasslands. The field work was carried out in the Považsky Inovec Mts (Western Carpathians) during 1993–1995. The permanent plot in the Poo badensis-Festucetum pallentis was located on a steep, strongly eroded S-facing slope covered with dolomite outcrops, scree and sparse vegetation (20%) dominated by Festuca pallens. The permanent plot in the Festuco pallentis-Caricetum humilis was located on the even ridge plateau with shallow stony soil and vegetation covering about 70% dominated by Carex humilis and Festuca pallens. In comparison to other grasses Festuca pallens had a very low rate of leaf turnover. The highest leaf birth rates and the lowest leaf death rates were observed in June. Leaf mortality was uniformly distributed in time without a distinct minimum or maximum. For the surviving tillers the leaf production exceeded the leaf mortality during the whole growing season. The steady net gain of leaves in tillers was not interrupted by the parallel process of tillering. Among the leaf cohorts the leaves produced in May had the longest leaf blades. Leaves grew during the whole year. The winter cold and summer drought might slow down the growth rate or interrupt the growth. The growth of a leaf blade took five to eight weeks. Leaf life span was estimated to 150–200 days (time from leaf appearance at the apex to the complete loss of its green assimilating parts). In comparison to other grasses Festuca pallens belongs to the species with the longest leaf life span. The effect of environmental factors on leaf demography was followed by the comparison of two populations belonging to two phytosociological associations differing mostly in habitat xericity. Differences were revealed in the following characteristics: length of leaf blade in cohorts born during May and June, maximum length of a leaf blade in a tiller and daily increments in May and June. The course of leaf natality and mortality was similar in the studied populations.     

五大连池新期火山熔岩台地3种共有植物的叶功能性状及其相互关系

以五大连池新期火山熔岩台地13个稳定的自然植物群落共有种香杨、万年蒿、岩败酱为研究对象,测定其叶干物质含量(LDMC)、比叶面积(SLA)、叶氮含量(LNC)、叶磷含量(LPC)、叶氮磷比(N/P)等叶功能性状,研究不同生境下3种植物叶功能性状的变异特征,探讨新期火山熔岩台地植物叶功能性状相互之间的内在联系及其对环境的适应性。结果表明:(1)3种植物LDMC变化在0.294~0.359g·g~(-1)之间,SLA变化在9.082~22.347m2·kg~(-1)之间;与其他区域的研究结果相比,新期火山熔岩台地植物的LDMC值相对偏大、SLA值相对偏小,说明五大连池新期火山熔岩台地的植物采用高LDMC低SLA策略适应贫瘠恶劣的环境。(2)3种植物LNC变化在9.690~15.710g·kg~(-1)之间,LPC变化在0.669~1.078g·kg~(-1)之间,叶钾含量(LKC)变化在10.410~29.830g·kg~(-1)之间,叶N/P的变化在9.781~20.990之间;与其他区域的研究结果相比,新期火山熔岩台地植物LNC和LPC值相对偏小,LKC和N/P值相对偏大,说明五大连池新期火山熔岩台地的植物生长主要受磷素的限制,而且LNC偏小和LKC偏高与新期火山熔岩台地土壤中全N、全K的含量变化特征有关。(3)通过ANVOA分析发现,群落间植物叶功能性状的变异幅度较小,3种植物之间N/P、LDMC和SLA的变异幅度也较小,这说明在水分和养分极缺的火山熔岩台地植物对其生境的适应对策相似。(4)Pearson相关分析发现,SLA与LDMC以及N/P与LDMC和SLA均呈显著负相关关系。     

西藏紫花针茅叶功能性状沿降水梯度的变化

植物叶功能性状与环境因子的关系是近10年来植物生态学的研究热点。该文以广泛分布于青藏高原干旱、半干旱草地的优势植物种紫花针茅(Stipa purpurea)为研究对象, 沿降水梯度(69-479 mm)系统测定了日土、改则、珠峰、当雄和纳木错5个调查地点紫花针茅比叶面积(SLA)、单位重量和单位面积叶氮含量(N_mass, N_area)、叶密度和厚度等叶功能性状以及土壤全氮含量等因子, 试图验证干旱胁迫地区同一物种内SLA-N_mass关系沿降水梯度的策略位移现象是否具有普遍性, 并对是否出现策略位移现象提出可能的解释。研究结果表明: 1) SLA和N_mass与生长季温度和降水以及土壤全氮含量均没有显著关系, SLA与N_mass的关系在干旱半干旱区(年降水/蒸发比< 0.11)与半湿润区(年降水/蒸发比> 0.11)之间并没有出现典型的位移现象; 2)叶密度是决定半湿润区SLA变化的主导因子, 而叶厚度则是干旱半干旱区SLA变化的控制因子, 两者与SLA均呈负相关, 随着温度增加或降水减少, 叶厚度增加而叶密度降低, 导致SLA随温度和降水变化不明显; 3)半湿润区的叶密度增加引起N_area增加, 而干旱半干旱区的叶厚度增加并没有造成N_area的显著变化, 导致N_area沿降水梯度没有显著变化; 4)紫花针茅地上生物量与N_area具有显著正相关关系, 表明N_area的增加有助于提高植被生产力。结果表明, 在干旱胁迫下, 植物通过增加叶厚度来维持不变的N_area可能有助于保持与较湿润地区相似的光合生产和水分利用效率。叶厚度和叶密度对比叶面积的相对影响在干旱半干旱区与半湿润区之间发生转变, 这为进一步检测高寒草地植被的水分限制阈值提供了新思路。