Leaf-shape variation of Paederia foetida in Japan: reexamination of the small, narrow leaf form from Miyajima Island

Variations in Paederia foetida L. leaf shape were examined to evaluate the taxonomic validity of the small, narrow leaf form of P. foetida f. microphylla Honda from Miyajima Island, Honshu, Japan. There is considerable variation in P. foetida individuals in terms of leaf size and leaf index (leaf length:leaf width ratio). On Miyajima Island, some individuals have narrow leaves with a high leaf index value, a phenotype represented by the type specimen of P. foetida f. microphylla, and some do not. Given that the leaf size of individuals from Miyajima Island is smaller than that of individuals from other localities in Japan, and that the small leaf phenotype is stable even under cultivation, P. foetida f. microphylla is classified as the form having the smallest leaf size. Anatomical examination of leaf blades revealed that the large variation in leaf size was attributable to variation in the number of leaf cells but not to differences in cell size or cell shape. Based on these results, we discuss the endemism of P. foetida f. microphylla.     

LeafAnalyser: a computational method for rapid and large-scale analyses of leaf shape variation

A comprehensive understanding of leaf shape is important in many investigations in plant biology. Techniques to assess variation in leaf shape are often time-consuming, labour-intensive and prohibited by complex calculation of large data sets. We have developed LeafAnalyser, software that uses image-processing techniques to greatly simplify the measurement of leaf shape variation. LeafAnalyser places a large number of evenly distributed landmarks along leaf margins and records the position of each automatically. We used LeafAnalyser to analyse the variation in 3000 leaves from 400 plants of Antirrhinum majus . We were able to summarise the major trends in leaf shape variation using a principal components (PC) analysis and assess the changes in size, width and tip-to-base asymmetry within our leaf library. We demonstrate how this information can be used to develop a model that describes the range and variation of leaf shape within standard wild-type lines, and illustrate the shape transformations that occur between leaf nodes. We also show that information from LeafAnalyser can be used to identify novel trends in shape variation, as low-variance PCs that only affect a subset of position landmarks. These results provide a high-throughput method to calculate leaf shape variation that allows a large number of leaves to be visualised in higher-dimensional phenotypic space. To illustrate the applicability of LeafAnalyser we also calculated the leaf shape variation in 300 leaves from Arabidopsis thaliana .     

The Morphological Variation and the Karyotypical Characters of Dendranthema indicum and D. lavandulifolium

Dendranthema indicum L. and D. lavandulifolium (Fisch. ex Trautv. ) Ling et Shih are two species which are greatly variable in morphology. The specimens collected from different habitats show distinct variation in characters such as leaf division、 leaf pubescence、 size of capitula and number of capitula. Some authors (Li 1983; Du 1989; Nakata 1987)have studied the chromosome numbers and karyotypes of D. indicum and D. lavandulifolium and reported the basic chromosome number and polyploidy of these two species. But some questions, such as the patterns and trends of morphological variation, the relationships between morphological and karyotypical variations and the phylogenetic relationship between D. indicum and D. lavandulifolium remain unclear . Based on wide field investigation and cultivation experiments, the present authors made a thorough study on D. indicum and D. lavandulifolium from morphology、 cytology with reference to geographical distribution and habit conditions. The results show that the amount of leaf pubescence of D. indicum is closely related to the water content in its habitat; the diameter of capitula decreases, while the number of capitula of D. lavandulifolium increases with the increase in elevation. Comparing the karyotypical constitution of different populations of D. indicum and D. lavandulifolium, the authors infer that natural hybridization, both intraspecific hybridization and hybridization between D. indicum and D. lavandulifolium, has played an important role in the origin of polyploids and in the evolutionary process of karyotypes of D. indicum and D. lavandulifolium. By a synthetic analysis from morphology, cytology and geography, the authors also suggest that D. indicum and D. lavandulifolium be sister species, derived from a common ancestor, and it is unreasonable to consider one of themevolved from the other.     

Quantitative variation within and among populations of Arabis serrata Fr. & Sav. (Brassicaceae)

Phenotypic and genetic variation within and among eight populations of Arabis serrata are documented in this study. This species shows great morphological variation throughout its geographical distribution in Japan. Plants are located in habitats with different types of soils and degree of disturbance. Half-sibs progenies from eight populations were collected and cultivated in a garden experiment. Nine morphological traits representing size and shape of rosette leaves were recorded. Univariate analyses of measured traits showed that phenotypic means differed among populations for all characters. Leaves of plants from disturbed habitats had the longest petioles (lanceolate) and plants from limestone habitats showed the most roundness in leaf shape (ovate). The northernmost populations always revealed the smallest leaves. Multivariate principal component analyses also showed that leaf shape and size varied among populations. The first three principal components explained 98.5% of the variation. Coefficients of variation had a very wide range and differed from one population to another. Some traits (e.g. leaf width/leaf length ratio) were consistently less variable while others (e.g. leaf area and petiole length) were more plastic. All traits had significant genetic variance in all populations. Intra-class correlation coefficients differed for most of the traits and each population presented a different range of values. Most of the leaf traits were intercorrelated in all the populations studied, although some populations were integrated more tightly for some traits. Populations of A. serrata are differentiated in phenotypic means but they display a mosaic of traits with slight morphological differences in each locality (i.e. a quantitative genetic variation). Some traits can be correlated to the habitats that they occupy but for some of them it is difficult to assign an actual adaptive value.     

Biosystematic Studies on Adenophora potaninii Korsh. Complex (Campanulaceae) III. Genetic Variation and Taxonomic Value of Morphological Characters

Adenophora potaninii Korsh．complex is a morphologically variable group including six species．As shown in many species in the genus Adenophora,the variability of many morphological characters in the complex under investigation is amazingly great,which brings difficulties in the delimitation and indentification of taxa．In the present study,six populations representing five species in the complex were systematically sampled to investigate the pattern of morphological variation within population and to detect the genetic basis of the variation with progeny tests and controlled crosses．The results are as follows： Great morphological variation within population is found (Fig.1),including characters such as leaf shape,teeth number and size of leaf margins,teeth number of calyx lobes,and indumentum on the surface of stems and leaves,which were previously considered as diagnotic．Systematic sampling and statistical analysis show that the differences in the above characters exhibit continuous patterns of variation within population (Fig．1,2),though they are highly correlated (Table 2)．From progeny testing and crossing between two contrast types of individuals (oblong,dentate and pubescent leaves vs．narrow,entire and glabrous leaves)it is evident that those characters show continuous variability in segregating progenies (Fig.3,4)and appear to be influenced by large numbers of loci with individually slight effects．Since most wild populations were highly heterozygous for those characters,the individuals in one extreme of variation can produce their variable offsprings including individuals similar to those in the other extreme (Fig. 3). This is also true for the indumentum density on stems and teeth number of calyx lobes. In this complex, A. biformifolia Y. Z. Zhao, A. bockiana Diels and A. polydentata P.F. Tu et G. J. Xu were described exclusively based on leaf shapes, teeth number of leaf margins and calyx lobes, teeth size of leaf margins, and indumentum on the surface of stemsand leaves. It is demonstrated, however, that the variation of those characters is of a quantitative nature and show no discontinuities. As a result, genetic analysis of diagnostic characters along with their sympatric distribution and same habitats, strongly suggest that A.biformifolia as well as A. bockiana and A. polydentata are actually the extreme individuals within A. wawreana and A. potaninii respectively and should not be recognized as taxa at any taxonomic level.     

Natural hybridization between a deciduous (Nothofagus antarctica, Nothofagaceae) and an evergreen (N. dombeyi) forest tree species: evidence from morphological and isoenzymatic traits

BACKGROUND AND AIMS: Trees with a partial leaf-shedding pattern and other morphological features a priori considered intermediate between those of the deciduous Nothofagus antarctica (G. Forster) Oersted and the evergreen N. dombeyi (Mirb.) Oersted (Nothofagaceae) were found in natural stands. The hybridization between a deciduous and an evergreen species of Nothofagus has not been reported so far in natural communities. METHODS: The putative hybrids and the two presumed parental species were compared using 14 enzyme systems as well as shoot, leaf and reproductive morphology. KEY RESULTS: Six enzyme systems showed good resolution (MDH-B, IDH, SKDH, 6-PGDH, GOT and PGI) and in four of them (PGI, MDH-B, SKDH and 6-PGDH) the putative hybrids showed intermediate zymogram patterns between N. antarctica and N. dombeyi. Both principal coordinates analysis on isozyme data and principal components analysis (PCA) on quantitative morphological traits of shoots and leaves separated both parental species and located the putative hybrids closer to N. antarctica than to N. dombeyi. In the PCA, the number of basal cataphylls and the length : width ratio of leaves were the variables most discriminating among shoots of the three entities. The putative hybrids were intermediate between both species regarding leaf vernation, outline and venation, variation in leaf shape (length/width) with position on the parent shoot and in staminate inflorescence and cupule morphology. For other morphological traits, the putative hybrids resembled one of the parental species or differed from both species (e.g. valve morphology). CONCLUSIONS: Isoenzymatic and morphological data sets support the idea of the hybrid nature (probably F1 generation) of the semi-deciduous trees found. Nothofagus antarctica and N. dombeyi are probably more closely related than previously assumed. The relevance of pollen type in revealing evolutionary relationships between Nothofagus species is supported, and that of leaf-shedding pattern is rejected.     

菊属与蒿属植物苗期抗蚜虫性鉴定

为发掘对蚜虫有抗性的菊花近缘种属植物,对11份菊属及蒿属植物苗期进行了田间和室内抗蚜虫性鉴定。结果发现:在田间鉴定条件下,随着平均蚜害指数(I*)值的升高,蚜虫抗性等级逐渐降低,其中:黄金艾蒿、黄蒿、香蒿的I*值为0.00,属于免疫性;牡蒿的I*值为0.13,属于高抗性;大岛野路菊的I*值为0.25,属于抗性;异色菊的I*值为0.37,属于中抗性;那贺川野菊、龙脑菊的I*值为0.51—0.75,属于感性;泡黄金菊、紫花野菊、菊花脑的I*值>0.75,属于高感性;在室内鉴定条件下,不同材料对蚜虫的抗性表现出很大差异,根据繁殖倍率的不同,11份材料分为5个抗性等级,其中黄金艾蒿、牡蒿、黄蒿、香蒿繁殖倍率为0.00,属于极高抗性;大岛野路菊繁殖倍率为5.86,属于中抗性;异色菊、那贺川野菊、龙脑菊繁殖倍率为7—10,属于低抗性;泡黄金菊、紫花野菊、菊花脑繁殖倍率>10,属于无抗性。综合分析表明:田间鉴定与室内鉴定结果基本一致,其中黄金艾蒿、黄蒿等8份材料田间和室内鉴定结果无差别;牡蒿田间鉴定表现为高抗性,而室内鉴定表现为极高抗性;大岛野路菊田间鉴定表现为抗性,而室内鉴定表现为中抗性;异色菊田间鉴定表现为中抗性,而室内鉴定表现为低抗性。黄金艾蒿、黄蒿、牡蒿、香蒿、大岛野路菊对蚜虫表现良好抗性,可用于栽培菊花抗蚜性遗传改良。     

Reproducibility of fluctuating asymmetry measurements in plants: Sources of variation and implications for study design

Fluctuating asymmetry (FA), i.e. small, non-directional deviations from perfect symmetry in morphological characters, increases under genetic and/or environmental stress. Ecological and evolutionary studies addressing FA became popular in past decades; however, their outcomes remain controversial. The discrepancies might be at least partly explained by inconsistent and non-standardised methodology. Our aim was to improve the methodology of these studies by identifying factors that affect the reproducibility of FA measurements in plant leaves. Six observers used a highly standardised measurement protocol to measure FA using the width, area and weight of the same set of leaves of 10 plant species that differed in leaf size, shape of the leaf margin and other leaf traits. On average, 24% of the total variation in the data was due to measurement error. Reproducibility of measurements varied with the shape of leaf margin, leaf size, the measured character and the experience of the observer. The lowest reproducibility of the width of leaf halves was found for simple leaves with serrate margins and the highest for simple leaves with entire margins and for compound pinnate leaves. The reproducibility was significantly lower for the weight of leaf halves than for either their width or area, especially for plants with small leaves. The reproducibility was also lower for measurements made by experienced observers than by naïve observers. The size of press-dried leaves decreased slightly but significantly relative to fresh leaves, but the FA of press-dried leaves adequately reflected the FA of fresh leaves. In contrast, preservation in 60% ethanol did not affect leaf size, but it decreased the width-based values of FA to 89.3% of the values measured from fresh leaves. We suggest that although reproducibility of leaf FA measurements depends upon many factors, the shape of the leaf margin is the most important source of variation. We recommend, whenever possible, choosing large-leaved plants with entire leaf margins as model objects for studies involving measurements of FA using the width of leaf halves. These measurements should be conducted with high accuracy from images of fresh or press-dried leaves.     

Correlated evolution of leaf shape and trichomes in Begonia dregei (Begoniaceae)

Structural features of leaves, including size, shape, and surfaces, vary greatly throughout the plant kingdom. In both functional and phylogenetic analyses of leaves, the various morphological aspects are often considered independently of each other, although it is likely that many combinations of features do not occur at random due to either functional constraint or genetic correlation. The distribution of variation in leaf morphology in the highly variable Begonia dregei species complex was examined in natural populations and in F(2) offspring from a cross between plants from two populations. Leaf shape was quantified using several morphometric measures, and trichomes on leaves were counted and measured. Correlations between leaf shape and the numbers and size of trichomes were examined. There were significant correlations between the shapes of leaves and the presence, number, and size of trichomes among populations and in hybrid plants. Deeply incised leaves had larger numbers of longer trichomes at the sinuses. Higher numbers of trichomes on upper leaf surfaces occurred together with trichomes at the petiole and on the abaxial surface. The potential for independent evolution of leaf shape and trichomes in this group is limited. Hypotheses to explain the correlated development of leaf shape and trichomes are discussed.     

Quantitative comparison of foliage development amongDryopteris monticola,D. tokyoensis and a putative hybrid,D. kominatoensis in Northern Japan

Developmental leaf architecture was quantitatively described in terms of measurements of various parameters on leaf blade from different size of sporophytes inDryopteris monticola, D. tokyoensis and a putative hybrid,D. kominatoensis in the natural site of Hokkaido, to compare the ontogenetic differentiation in foliage structure among allied ferns. The morphological stage of leaf and sporophyte was tentatively quantified by the number of midrib branches of the leaf (NV, number of veins), which exhibited a significant correlation to the leaf-shape complexity from a circle (DI=marginal length/2×(3.14×square)^1/2) of leaf blade. D. kominatoensis showed intermediate values between others in following characters; DI increase, maximum NV (also blade length), maximum number of costa branches of pinnae (NVMP), number of costa branches of the lowest pinna (NVLP), difference between NVMP and NVLP (NVMP-NVLP), during heteroblastic leaf development. A larger number of leaves per sporophyte was found inD. kominatoensis than in others. The fertility rate (%) and initiation of fertility (IF) in the relative developmental stage (RDS) ofD. kominatoensis shifted to that ofD. tokyoensis, while the order of pinnae with NVMP shifted to that ofD. monticola. Even in the intermediate characters inD. kominatoensis, slight shifts in characters to those of putative parents were found during heteroblastic leaf development. Contribution No. 3145 from the Institute of Low Temperature Science, Hokkaido University.     

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.     

Phylogenetic Relationship of Dendranthema (DC.) Des Moul. Revealed by Fluorescent In Situ Hybridization

Phylogenetic relationships of the different species in the genus Dendranthema (DC.) Des Moul. were estimated based on chromosome fluorescent in situ hybridization (FISH) with 18S-26S rDNA of Arabidopsis and genomic DNA of Dendranthema as probes. The results revealed that there was no positive correlation between the number of nuclear organization region (NOR) loci and the ploidy of Dendranthema.The exact cytogenetic information of NORs about 14 operational taxonomic units (OTUs) indicated that D.vestitum (Hemsl.) Ling et Shih was closer to the cultivars than other putative species, whereas D. zawadskii (Herb.) Tzvel. was the most distinct. The ambiguously distributed signals of genomic in situ hybridization (GISH) with genomic DNA of lower ploidy species as probes suggested that different genomes among Dendranthema were mixed. The result also indicated the limitation of GISH in studies on the phylogenetic relationships of the different species in this genus Dendranthema and on the origin of cultivated chrysanthemums. Based on these results and previous research, the origin of Chinese cultivated chrysanthemum is discussed.     

Organ shape and size: a lesson from studies of leaf morphogenesis

Control of the shape and size of indeterminate organs, such as roots and stems, is directly related to the control of the shape and size of the cells in these organs, as predicted by orthodox cell theory. For example, the polarity-dependent growth of leaf cells directly affects the polar expansion of leaves. Thus, the control of leaf shape is related to the control of the shape of cells within the leaf, as suggested by cell theory. By contrast, in determinate organs, such as leaves, the number of cells does not necessarily reflect organ shape or size. Genetic evidence shows that a compensatory system(s) is involved in leaf morphogenesis, and that an increase in cell volume can be triggered by a decrease in cell number and vice versa. Studies of chimeric leaves also suggest interaction between leaf cells that coordinates the behaviour of these cells at the organ level. Moreover, leaf size also appears to be coordinated at the whole-plant level. The recently hypothesised neo cell theory describes how leaf shape- and size-control mechanisms control leaf shape at the organ-level via cell-cell interaction.     

Effect of Salinity on the Composition, Number and Size of Epidermal Cells along the Mature Blade of Wheat Leaves

Salinity inhibits leaf growth in association with changes in cell size. The objective of this study was to determine the spatial distributions of the composition, number and dimensions of epidermal cells in the mature blades of leaf four of wheat seedlings under saline conditions. Plants were grown in loamy soil either with or without 120 mmol/L NaCl in a growth chamber, and harvested after leaf four was fully developed. The results of the spatial distribution analyses of width along the blade showed that salinity not only reduced the width of the leaf blade, but that it also altered the distribution pattern of blade width along the leaf axis. The reduction in the final size of the leaf blade was associated with a reduction in the total number of epidermal cells and in their widths and lengths. This study also revealed the spatial effects of salinity on the blade and epidermal cell dimensions along the leaf axis. In particular, salinity inhibited the total cell number for interstomatal, sister and elongated cells, implying that cell division in wheat leaves is inhibited by salinity. However, the lengths of interstomatal cells were not affected by salinity （unlike those for the sister and elongated cells）, suggesting the relative contributions of cell length and numbers to the reduction in the final length of the blade under salinity is dependent on cell type.     

Phylogenetic Relationship of Dendranthema （DC.） Des Moul. Revealed by Fluorescent In Situ Hybridization

Phylogenetic relationships of the different species in the genus Dendranthema (DC.) Des Moul. were estimated based on chromosome fluorescent in situ hybridization (FISH) with 18S-26S rDNA of Arabidopsis and genomic DNA of Dendranthema as probes. The results revealed that there was no positive correlation between the number of nuclear organization region (NOR) loci and the ploidy of Dendranthema.The exact cytogenetic information of NORs about 14 operational taxonomic units (OTUs) indicated that D.vestitum (Hemsl.) Ling et Shih was closer to the cultivars than other putative species, whereas D. zawadskii (Herb.) Tzvel. was the most distinct. The ambiguously distributed signals of genomic in situ hybridization (GISH) with genomic DNA of lower ploidy species as probes suggested that different genomes among Dendranthema were mixed. The result also indicated the limitation of GISH in studies on the phylogenetic relationships of the different species in this genus Dendranthema and on the origin of cultivated chrysanthemums. Based on these results and previous research, the origin of Chinese cultivated chrysanthemum is discussed.     

Heteroblasty in the moss,Aphanoregma patens (Physcomitrella patens), results from progressive modulation of a single fundamental leaf developmental programme

Abstract

Leaves at the apex of a mature Aphanoregma patens (Hedw.) Lindb. (Physcomitrella patens (Hedw.) Bruch Schimp. in B.S.G.) gametophore differ markedly in size and form from those at its base. To determine how these differences are produced during development, we first examined qualitative and quantitative differences between successive leaves along the stem and among leaves at different developmental stages. Differences between successive leaves were slight and cumulative. Local changes in cell number and size combined to produce a regularly shaped and approximately bilaterally symmetrical leaf suggesting that cell division and cell expansion are regionally regulated and coordinated at the organ level. The midrib and marginal teeth are discrete characters, which were prefigured by changes in cell shape in leaves that lacked these characters. In leaf primordia, cell proliferation was responsible for most of the changes in leaf form and size early in development and may have continued as cell expansion took over as the primary contributor to leaf growth and morphogenesis. Thus, leaf heteroblasty in Physcomitrella probably results from modulation of a single developmental programme by external and/or internal forces, which alter progressively in intensity as a gametophore grows. We applied exogenous cytokinin and auxin separately to growing cultures to explore their effects on leaf growth. Cytokinin and auxin stimulated leaf cell division and leaf cell elongation, respectively. Also, young upper leaves of gametophores exposed to exogenous auxin closely resembled basal leaves of untreated plants. Therefore, endogenous cytokinins and auxins may be among the modulating internal forces involved in leaf morphogenesis and the establishment of leaf heteroblasty.     

泡沙参复合体(桔梗科)的物种生物学研究——Ⅲ.性状的遗传变异及其分类价值

阐明性状变异的遗传基础是评估性状分类价值以及揭示类群关系和进化过程的前提。本文根据泡沙参复合体内6个天然居群的材料,利用居群样本分析、子代测定和杂交试验等手段,结合统计学方法对该复合体一些在以往类群划分中起鉴别作用的形态性状进行了遗传学分析。结果表明,叶形、叶缘锯齿数目和大小、茎叶被毛、花萼裂片齿数、花梗长短等性状都是遗传性比较强的性状,而且性状之间有很强的相关性。但是,这些性状在居群内呈现连续变异且变异幅度非常大,以致于变异的两个极端个体被分别作为不同的种处理。通过对野外居群样本的统计分析和对极端变异类型个体所进行的子代测定及其人工杂交,进一步证实上述性状是受多基因决定的数量性状,其变异幅度大,变异式样中并不存在任何程度的间断性,因而至少在本复合体内没有鉴别意义。因此,以往建立在上述性状变异基础上的3个类群(A.biformifolia Y.Z.Zhao;A.bockiana Diels和A.polydentata P.F.Tu et G.I.Xu)均不成立。最后,本文对形态性状变异的遗传基础及其与类群划分的关系进行了讨论。