Comparative phylogeography of two closely related <Emphasis Type="Italic">Viola</Emphasis> species occurring in contrasting habitats in the Japanese archipelago

Similar distribution ranges shared by closely related plant species may have been shaped through different migration histories if those species have differing habitat preference. To test this hypothesis, phylogeographical patterns and population genetic structures were compared between two sister Viola species: Viola eizanensis preferring woodland and V. chaerophylloides var. sieboldiana preferring grassland, both being native to the Japanese Archipelago. Amplified fragment length polymorphism (AFLP) was used for phylogenetic reconstruction, together with Bayesian ancestry analysis, AMOVA, analysis of genetic diversity statistics, and analysis of the relative contribution of each population to total diversity. The results indicated that V. eizanensis had two distinct lineages occurring in the western and eastern part of Japan, but such lineages are not distinct in V. chaerophylloides var. sieboldiana. Both species exhibited the low genetic diversity and high between-population differentiation typical of selfing plants. In V. chaerophylloides var. sieboldiana, one particular population made a significantly higher contribution to the total heterozygosity (H _T), whereas in V. eizanensis, no population was identified as making a particularly higher contribution to H _T. These findings suggest that V. eizanensis had been isolated in two large glacial refugia, whereas populations of V. chaerophylloides var. sieboldiana were restricted to a single small refuge. Different light requirements between these two closely related species probably caused these differing responses to climatic change during the last ice age. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Constitutive knox1 gene expression in dandelion (Taraxacum officinale, Web.) changes leaf morphology from simple to compound

Seed plants with compound leaves constitute a polyphyletic group, but studies of diverse taxa show that genes of the class 1 KNOTTED-LIKE HOMEOBOX (KNOX1) family are often involved in compound leaf development. This suggests that knox1 genes have been recruited on multiple occasions during angiosperm evolution (Bharathan et al. in Science 296:1858–1860, 2002). In agreement with this, we demonstrate that the simple leaf of dandelion (Taraxacum officinale Web.) can be converted into a compound leaf by the constitutive expression of heterologous knox1 genes. Dandelion is a rosette plant of the family Asteraceae, characterised by simple leaves with deeply lobed margins and endogenous knox1 gene expression. Transgenic dandelion plants constitutively expressing the barley (Hordeum vulgare L.) hooded gene (bkn3, barley knox3) or the related bkn1 gene, developed compound leaves featuring epiphyllous rosettes. We discuss these results in the context of two current models of compound leaf formation.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Lachemilla jaramilloi and L. talamanquensis spp. nov. (Rosaceae) from Ecuador and Costa Rica

Two new species of Lachemilla (Rosaceae) are described and illustrated. Lachemilla jaramilloi Romoleroux &Morales‐Briones from Ecuador is characterized by its decumbent habit, villous‐hirsute pubescence, long basal petiole, and deeply tri‐parted leaf blades. The flowers are sessile to subsessile and subtended by episepals, and the hypanthia elongated and glabrescent. Lachemilla talamanquensis Romoleroux & Morales‐Briones from Costa Rica is distinguished by its sericeous‐villous, ascending branches, sub‐sessile distal leaves, tri‐lobed leaf blades, forming whorls with the stipule. The flowers are solitary or arranged in few‐flowered inflorescences, with big, dark, villous hypanthia, containing several achenes.     

Two new stoloniferous species of Viola (Violaceae) from China

Two new stoloniferous species of Viola (Violaceae) from southern China are described and illustrated. Viola nitida is recognizably different from V. fargesii as the plant is evergreen and glabrous throughout, the leaf blade adaxially nitid, base cuneate or shallowly cordate, and margin serrate. Viola maoershanensis is different from V. diffusa as the leaf blade is serrate, base cordate and not decurrent to petiole, the petiole wingless, the flowers larger, the petals bluish violet or pinkish white, and the lower petal obtuse at the apex. The chromosome numbers of the two new species were counted as 2n = 24 (V. nitida) and 2n = 44 (V. maoershanensis). The taxonomic positions of the two species are discussed. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 349–356.     

Effects of Leaf Shape and Boundary Layer Thickness on Photosynthesis in Cotton (Gossypium hirsutum)

In upland cotton (Gossypium hirsutum L.) certain varieties are available with the mutant character “okra” leaves. These deeply lobed leaves were found to have thinner boundary layers than their normal analogues. Apparent photosynthesis and transpiration measurements were made in field-grown stands under a variety of light intensities and carbon dioxide levels to assess the effect of leaf boundary layer diffusion resistance on photosynthetic efficiency. The thinner boundary layers associated with deeply lobed “okra” cotton failed to euhance carbon fixation rates per unit land area. It was concluded that the leaf boundary layer resistance under field conditions is small compared with the total CO₂ diffusion resistance.     

Ranunculus giordanoi sp. nov. from the R. auricomus complex (Ranunculaceae), central Apennines (Italy)

A new species, Ranunculus giordanoi, belonging to the R. auricomus complex is described from Latium, central Apennines (Italy). It grows in damp meadows on Monti della Laga (National Park of Gran Sasso and Monti della Laga), the largest sandstone massif of central Italy. The new species differs from the closely related R. silanus, endemic to Sila (Calabria, southern Apennines), by leaf aperture, shape of the basal leaves, entire or sometimes deeply lobed cauline leaves, gynoclinium without intervallum and longer carpellophores.     

CLAUSA restricts tomato leaf morphogenesis and GOBLET expression

Leaf morphogenesis and differentiation are highly flexible processes. The development of compound leaves is characterized by an extended morphogenesis stage compared with that of simple leaves. The tomato mutant clausa (clau) possesses extremely elaborate compound leaves. Here we show that this elaboration is generated by further extension of the morphogenetic window, partly via the activity of ectopic meristems present on clau leaves. Further, we propose that CLAU might negatively affect expression of the NAM/CUC gene GOBLET (GOB), an important modulator of compound‐leaf development, as GOB expression is elevated in clau mutants and reducing GOB expression suppresses the clau phenotype. Expression of GOB is also elevated in the compound leaf mutant lyrate (lyr), and the remarkable enhancement of the clau phenotype by lyr suggests that clau and lyr affect leaf development and GOB in different pathways.     

Geographical variation and genetic analysis of leaf shape inCrepis tectorum (Asteraceae)

Canonical variate analysis of plants raised in a uniform environment was used to study the pattern of geographical variation in leaf shape ofCrepis tectorum (Asteraceae). The diversity in leaf shape was much greater among populations confined to areas with exposed bedrock in the Baltic region than among weed populations scattered throughout Europe and Canada. A Ward's clustering linked outcrop populations from the archipelago of SW. Finland and the islands of Öland (Sweden) and Saaremaa (Estonia) due to the deeply lobed leaves characterizing these populations, while outcrop populations along the coast of E. Sweden were grouped due to their weakly lobed, narrow and dentate leaves. Most of the weed populations were grouped together but there was no tendency for the variation in this group to be related to habitat or geographical location. A mosaic of variation reflected in sharp (random) differentiation among local populations was superimposed on the large-scale ecogeographical pattern.—Crossing data indicated that most of the variation in degree of leaf dissection is governed by one major gene with deeply lobed leaves dominant over weakly lobed leaves. I suggest that the simple pattern of inheritance may have favoured rapid evolutionary changes in leaf shape, particularly in the Baltic area which emerged relatively late from the sea. Genetic correlations may have constrained the pattern of variation at higher taxonomic levels, since some of the trait associations detected in a segregating F₂ generation were also found at the among-population level.     

Ranunculus uttaranchalensis (Ranunculaceae): a new species from Western Himalaya,India

A new species of the genus Ranunculus, R. uttaranchalensis, is described from Gangotri National Park, Uttaranchal in Western Himalaya, India. It is distinguished from the closely allied R. lobatus in having radical leaves sub‐cordate with hairy lamina base and petiole hairy above; cauline leaves sessile, 3–7 lobed with linear to narrowly lanceolate lobes; sepals usually reddish brown, externally hairy; petals rounded‐obovate and oblong, receptacle glabrous except 1–3 hairs at the top.     

Developmental events leading to peltate leaf structure in <Emphasis Type="Italic">Tropaeolum majus</Emphasis> (Tropaeolaceae) are associated with expression domain changes of a YABBY gene

Peltate leaf architecture has evolved from conventional bifacial leaves many times in flowering plant evolution. Characteristics of peltate leaves, such as the differentiation of a cross zone and of a radially symmetric, margin-less petiole, have also been observed in mutants of genes responsible for adaxial-abaxial polarity establishment. This suggests that altered regulation of such genes provided a mechanism for the evolution of peltate leaf structure. Here, we show that evolution of leaf peltation in Tropaeolum majus, a species distantly related to Arabidopsis thaliana, was associated with altered expression of Tropaeolum majus FILAMENTOUS FLOWER (TmFIL), a gene conferring abaxial identity. In situ hybridization indicates that adaxial and abaxial domains are established in early leaf primordia as in species with bifacial leaves. Upon initiation of the cross zone by fusion of the blade margins, localized expansion of TmFIL to the upper leaf side could be seen, indicating a local loss of adaxial leaf identity. The observed changes in expression are consistent with a role of TmFIL in radialization of the petiole and circularization of the leaf blade margin by the cross zone. In addition, expression was observed in segment primordia and during expansion of the bifacial blade, suggesting additional roles for TmFIL in leaf development.     

优良单株家系辣木叶的表型性状分析

为挖掘辣木(Moranga oleifera)优良种质资源,对30个优良单株家系的叶片表型性状进行研究。结果表明,除叶形外,辣木不同家系间的叶柄和叶片颜色、复叶数、复叶柄长度和直径、复叶间距、叶长、叶宽均存在不同程度的差异。复叶数与复叶柄长度和直径、复叶间距、叶长、叶宽呈极显著正相关;主成分分析表明,叶长、叶宽、复叶柄长度和直径、复叶间距、叶柄和叶片颜色是区分辣木不同家系最主要的叶片性状指标。聚类分析结果表明,30个辣木家系可分为3大类,叶片表型性状存在显著差异的家系的遗传距离较远。因此,叶柄和叶片颜色、复叶数、复叶柄长度和直径、复叶间距、叶长、叶宽将为直观区分辣木家系提供参考。     

Leaf shape variation and herbivore consumption and performance: a case study with Ipomoea hederacea and three generalists

The effect of leaf shape variation on plant-herbivore interactions has primarily been studied from the perspective of host seeking behavior. Yet for leaf shape to affect plant-herbivore coevolution, there must be reciprocal effects of leaf shape variation on herbivore consumption and performance. We investigated whether alternative leaf morphs affected the performance of three generalist insect herbivores by taking advantage of a genetic polymorphism and developmental plasticity in leaf shape in the Ivyleaf morning glory, Ipomoea hederacea. Across four experiments, we found variable support for an effect of leaf shape genotype on insects. For cabbage loopers (Trichoplusia ni) and corn earworms (Helicoverpa zea) we found opposing, non-significant trends: T. ni gained more biomass on lobed genotypes, while H. zea gained more biomass on heart-shaped genotypes. For army beetworms (Spodoptera exigua), the effects of leaf shape genotype differed depending on the age of the plants and photoperiod of growing conditions. Caterpillars feeding on tissue from older plants (95 days) grown under long day photoperiods had significantly greater consumption, dry biomass, and digestive efficiency on lobed genotypes. In contrast, there were no significant differences between heart-shaped and lobed genotypes for caterpillars feeding on tissue from younger plants (50 days) grown under short day photoperiods. For plants grown under short days, we found that S. exigua consumed significantly less leaf area when feeding on mature leaves than juvenile leaves, regardless of leaf shape genotype. Taken together, our results suggest that the effects of leaf shape variation on insect performance are likely to vary between insect species, growth conditions of the plant, and the developmental stage and age of leaves sampled. Handling editor: May Berenbaum.     

<Emphasis Type="Italic">KNOX</Emphasis> overexpression in transgenic <Emphasis Type="Italic">Kohleria</Emphasis> (Gesneriaceae) prolongs the activity of proximal leaf blastozones and drastically alters segment fate

KNOX (knotted1-like homeobox) genes have a widely conserved role in the generation of dissected leaves. Ectopic KNOX activity in leaves in various angiosperm lineages causes leaf form changes that can elucidate how the configuration of leaf development evolved. We present an analysis of leaf morphology and morphogenesis in transgenic Kohleria lines overexpressing a heterologous KNOX gene. Kohleria, like many members of Gesneriaceae, has simple-serrated leaves with pinnate venation. KNOX overexpression causes prolonged segment proliferation in proximal, but not distal, parts of leaf blades. Elaborate dissected segments reiterate the zonation of the whole leaf, with organogenic activity persisting between a distal maturation zone and a proximal intercalary elongation zone. The architecture of vascular bundles is severely altered, with a reduced midvein and a more palmate venation. The initial establishment of organogenically competent primordial margins (marginal blastozones) and the onset of tissue differentiation in early stages of leaf development were similar in wild-type and KNOX overexpressing lines. However, leaves overexpressing KNOX often failed to fully mature, and persistent marginal blastozones were found at the base of blades in mature portions of the shoot. We conclude that KNOX-mediated perpetuation of marginal blastozones in Kohleria is sufficient to induce a set of processes that result in highly dissected leaflets, which are unusual in this plant family. Spatial confinement of blastozones between an early maturing tip and a late elongating petiole zone reflects the presence of distinct maturation processes that limit the ability of the leaf margins to respond to ectopic KNOX gene expression.     

Functional Analysis and RNA Sequencing Indicate the Regulatory Role of Argonaute1 in Tomato Compound Leaf Development

Regardless of whether a leaf is simple or compound, the mechanism underlying its development will give rise to a full comprehension of plant morphogenesis. The role of Argonaute1 (AGO1) in the development of simple leaves has been established, but its role in the development of compound leaves remains to be characterized. In this paper, a virus-induced gene silencing (VIGS) strategy was used to dramatically down-regulate the expression of AGO1 ortholog in tomatoes, a model plant for research into compound leaves. AGO1-silenced tomato compound leaves exhibited morphological defects of leaf adaxial-abaxial and trichome development. Analysis of global gene expression profiles indicated that the silencing of AGO1 in tomato compound leaf caused significant changes in the expression of several critical genes, including Auxin Response Factor 4 (ARF4) and Non-expressor of PR5 (NPR5), which were involved in adaxial-abaxial formation and IAA15 that was found to contribute to growth of trichomes as well as Gibberellic Acid Insensitive (GAI) which participated in hormone regulation. Collectively, these results shed light on the complicated mechanism by which AGO1 regulates compound leaf development.     

Abnormal cell divisions in leaf primordia caused by the expression of the rice homeobox geneOSH1 lead to altered morphology of leaves in transgenic tobacco

Transgenic tobacco plants were generated carrying a rice homeobox gene,OSH1, controlled by the promoter of a gene encoding a tobacco pathogenesis-related protein (PR1a). These lines were morphologically abnormal, with wrinkled and/or lobed leaves. Histological analysis of shoot apex primordia indicated arrest of lateral leaf blade expansion, often resulting in asymmetric and anisotropic growth of leaf blades. Other notable abnormalities included abnormal or arrested development of leaf lateral veins. Interestingly,OSH1 expression was undetectable in mature leaves with the aberrant morphological features. Thus,OSH1 expression in mature leaves is not necessary for abnormal leaf development. Northern blot and in situ hybridization analyses indicate thatPR1a-OSH1 is expressed only in the shoot apical meristem and in very young leaf primordia. Therefore, the aberrant morphological features are an indirect consequence of ectopicOSH1 gene expression. The only abnormality observed in tissues expressing the transgene was periclinal (rather than anticlinal) division in mesophyll cells during leaf blade initiation. This generates thicker leaf blades and disrupts the mesophyll cell layers, from which vascular tissues differentiate. TheOSH1 product appears to affect the mechanism controlling the orientation of the plane of cell division, resulting in abnormal periclinal division of mesophyll cell, which in turn results in the gross morphological abnormalities observed in the transgenic lines.     

Mechanisms for minimizing height‐related stomatal conductance declines in tall vines

The ability to transport water through tall stems hydraulically limits stomatal conductance (g_s), thereby constraining photosynthesis and growth. However, some plants are able to minimize this height‐related decrease in g_s, regardless of path length. We hypothesized that kudzu (Pueraria lobata) prevents strong declines in g_s with height through appreciable structural and hydraulic compensative alterations. We observed only a 12% decline in maximum g_s along 15‐m‐long stems and were able to model this empirical trend. Increasing resistance with transport distance was not compensated by increasing sapwood‐to‐leaf‐area ratio. Compensating for increasing leaf area by adjusting the driving force would require water potential reaching ?1.9 MPa, far below the wilting point (?1.2 MPa). The negative effect of stem length was compensated for by decreasing petiole hydraulic resistance and by increasing stem sapwood area and water storage, with capacitive discharge representing 8–12% of the water flux. In addition, large lateral (petiole, leaves) relative to axial hydraulic resistance helped improve water flow distribution to top leaves. These results indicate that g_s of distal leaves can be similar to that of basal leaves, provided that resistance is highest in petioles, and sufficient amounts of water storage can be used to subsidize the transpiration stream.     

DISPERSAL ECOLOGY OF VANCOUVERIA (BERBERIDACEAE)

All Vancouveria species are habitual myrmecochores of the Viola odorata type. The anatropous, bitegmic seed develops an unusual elaiosome in the form of a large, empty, folded and lobed sac of epidermal tissue. The thin-walled assimilating capsule dehisces along an oblique weak zone that probably represents the suture between two fused and unequal carpels. In V. hexandra the capsule opens early to expose immature assimilating seeds. Decurved pedicels, down-turned fruits, and weak funiculi produce tachyspory. The related Epimedium, Jeffersonia, and Plagiorhegma have appendaged seeds and other possible myrmecochorous features resembling those of Vancouveria. Myrmecochory was probably established in the early or middle Tertiary period. The Vancouveria-Epimedium complex indicates that enormous plant migrations may be accomplished by means of ants.