Altered expression patterns of TCP and MYB genes relating to the floral developmental transition from initial zygomorphy to actinomorphy in Bournea (Gesneriaceae)

The shift from zygomorphy to actinomorphy has been intensively studied in molecular genetic model organisms. However, it is still a key challenge to explain the great morphological diversity of derived actinomorphy in angiosperms, since different underlying mechanisms may be responsible for similar external morphologies. Bournea (Gesneriaceae) is of particular interest in addressing this question, as it is a representative of primarily derived actinomorphy characteristic of a unique developmental transition from zygomorphy to actinomorphic flowers at anthesis. Using RNA in situ hybridization, the expression patterns were investigated of three different Bournea orthologues of TCP and MYB genes that have been shown to control floral symmetry in model species. Here, it is shown that the initial zygomorphic pattern in Bournea is likely a residual zygomorphy resulting from conserved expression of the adaxial (dorsal) identity gene BlCYC1. As a key novel event, the late downregulation of BlCYC1 and BlRAD and the correlative changes in the late specific expression of the abaxial (ventral) identity gene BlDIV should be responsible for the origin of the derived actinomorphy in Bournea. These results further indicate that there might be diverse pathways in the origin and evolution of derived actinomorphy through modifications of pre-existing zygomorphic developmental programs under dynamics of regulatory networks.     

花对称性的研究进展

花对称性(floral symmetry)是被子植物花部结构的典型特性之一,主要有辐射对称和两侧对称两种形式。被子植物初始起源的花为辐射对称,而两侧对称的花则是由辐射对称的花演变而来。两侧对称的花部结构是被子植物进化过程中的一个关键的革新,被认为是物种形成和分化的关键推动力之一。近年来有关花对称性的形成和进化机制的研究在植物学科的不同领域均取得了长足的进展。本文综述了花对称性在发育生物学、传粉生物学、生殖生态学及分子生物学等方面的研究进展。两侧对称形成于被子植物花器官发育的起始阶段,随后贯穿整个花器官发育过程或者出现在花器官发育后期的不同阶段。花器官发育过程中一种或多种类型器官的败育以及特异性花器官结构的形成是两侧对称形成的主要原因。研究表明,在传粉过程的不同阶段,花对称性均会受到传粉昆虫介导的选择作用。相比辐射对称的花,两侧对称的花提高了特异性传粉者的选择作用,增加了花粉落置的精确性,进而确保了其生殖成功。花对称性的分子机理已经在多种双子叶植物中进行了深入的研究。现有的证据表明,CYC同源基因在花对称性的分子调控方面起着非常重要的作用。花对称性在被子植物进化过程中是如何起源,与其他花部构成之间是否协同作用,一些不符合一般模式的科属其花对称性的形成机制等都是今后要进一步研究的命题。     

The floral development and floral anatomy ofChrysosplenium alternifolium, an unusal member of the Saxifragaceae

The floral development and anatomy ofChrysosplenium alternifolium were studied with the scanning electron microscope and light microscope to understand the initiation sequence of the floral organs and the morphology of the flower, and to find suitable floral characters to interpret the systematic position of the genus within the Saxifragaceae. The tetramerous flower shows a highly variable initiation sequence. The median sepals and first stamens arise in a paired sequence resembling a dimerous arrangement, but the first sepal and stamen arise on the side opposite to the bract. Transversal sepals and stamens emerge sequentially, as one side often precedes the other; sepals and stamens occasionally arise on common primordia. Initiation of the gynoecium is more constant with two median carpel primordia arising on a sunken floral apex. Several flowers were found to be pentamerous with a 2/5 initiation sequence. Flowers were invariably found to be apetalous without traces of petals in primordial stages; this condition is interpreted as an apomorphy. It is postulated that the development of a broad gynoecial nectary is responsible for the occurrence of an obdiplostemonous androecium. The gynoecium shows a number of anatomical particularities not observed in other Saxifragaceae. The presence and distribution of colleters is discussed.     

Development of the gametophytes, flower, and floral vasculature in Dichorisandra thyrsiflora (Commelinaceae)

The flowers of Dichorisandra thyrsiflora (Commelinaceae) are monosymmetric and composed of three sepals, three petals, six stamens, and three connate carpels. The anthers are poricidal and possess a wall of five cell layers (tapetum included). This type of anther wall, not previously observed in the Commelinaceae, is developmentally derived from the monocotyledonous type via an additional periclinal division and the persistence of the middle layers through anther dehiscence. Secondary endothecial thickenings develop in the cells of the two middle layers only. The tapetum is periplasmodial and contains raphides. Microsporogenesis is successive and yields both decussate and isobilateral tetrads. Pollen is shed as single binucleate grains. The gynoecium is differentiated into a globose ovary, hollow elongate style, and trilobed papillate stigma. Each locule contains six to eight hemianatropous to slightly campylotropous crassinucellar ovules with axile (submarginal) placentation. The ovules are bitegmic with a slightly zig-zag micropyle. Megagametophyte development is of the Polygonum type. The mature megagametophyte consists of an egg apparatus and fusion nucleus; the antipodals having degenerated. The floral vasculature is organized into an outer and inner system of bundles in the pedicel. The outer system becomes ventral carpellary bundles. All other floral vascular traces originate from the inner system.     

The development of symmetry,rigidity and anchorage in the structural root system of conifers

The stability of shallowly rooted trees can be strongly influenced by the symmetry of the ‘structural’ system of woody roots. Root systems of forest trees are often markedly asymmetric, and many of the factors affecting symmetry, including root initiation and the growth of primary and woody roots, are poorly understood. The internal and environmental factors that control the development, with respect to symmetry and rigidity, of shallow structural root systems are reviewed and discussed with particular reference to Sitka spruce (Picea sitchensis Bong. Carr.). Areas where there is insufficient knowledge are highlighted. A scheme is proposed that represents the root system as a set of spokes that are variable in number, size and radial distribution. Rigidity can vary between and along each of the spokes. The root system is presented as a zone of competition for assimilates, where allocation to individual roots depends upon their position and local variations in conditions. Factors considered include the production of root primordia of different sizes, effects of soil conditions such as the supply of mineral nutrients and water on growth of primary and woody roots, and the effect of forces caused by wind action on growth of the cambium, giving rise to roots which, in cross section, resemble I- or T-beams, and efficiently resist bending. This revised version was published online in June 2006 with corrections to the Cover Date.     

Analysis of PEAM4, the pea AP1 functional homologue, supports a model for AP1-like genes controlling both floral meristem and floral organ identity in different plant species

APETALA1 (AP1) and its homologue SQUAMOSA (SQUA) are key regulatory genes specifying floral meristem identity in the model plants Arabidopsis and Antirrhinum. Despite many similarities in their sequence, expression and functions, only AP1 appears to have the additional role of specifying sepal and petal identity. No true AP1/SQUA-functional homologues from any other plant species have been functionally studied in detail, therefore the question of how the different functions of AP1-like genes are conserved between species has not been addressed. We have isolated and characterized PEAM4, the AP1/SQUA-functional homologue from pea, a plant with a different floral morphology and inflorescence architecture to that of Arabidopsis or Antirrhinum. PEAM4 encodes for a polypeptide 76% identical to AP1, but lacks the C-terminal prenylation motif, common to AP1 and SQUA, that has been suggested to control the activity of AP1. Nevertheless, constitutive expression of PEAM4 caused early flowering in tobacco and Arabidopsis. In Arabidopsis, PEAM4 also caused inflorescence-to-flower transformations similar to constitutive AP1 expression, and was able to rescue the floral organ defects of the strong ap1-1 mutant. Our results suggest that the control of both floral meristem and floral organ identity by AP1 is not restricted to Arabidopsis, but is extended to species with diverse floral morphologies, such as pea.     

Defective cell proliferation in the floral meristem of alloplasmic plants of Nicotiana tabacum leads to abnormal floral organ development and male sterility

Flowers of an alloplasmic male-sterile tobacco line, comprised of the nuclear genome of Nicotiana tabacum and the cytoplasm of Nicotiana repanda, develop short, poorly-pigmented petals and abnormal sterile stamens that often are fused with the carpel wall. The development of flower organ primordia and establishment of boundaries between the different zones in the floral meristem were investigated by performing expression analysis of the tobacco orthologs of the organ identity genes GLO, AG and DEF. These studies support the conclusion that boundary formation was impaired between the organs produced in whorls 3 and 4 resulting in partial fusions between anthers and carpels. According to the investigations cell divisions and floral meristem size in the alloplasmic line were drastically reduced in comparison with the male-fertile tobacco line. The reduction in cell divisions leads to a discrepancy between cell number and cell determination at the stage when petal and stamen primordia should be initiated. At the same stage expression of the homeotic genes was delayed in comparison with the male-fertile line. However, the abnormal organ development was not due to a failure in the spatial expression of the organ identity genes. Instead the aberrant development in the floral organs of whorls 2, 3 and 4 appears to be caused by deficient floral meristem development at an earlier stage. Furthermore, defects in cell proliferation in the floral meristem of the alloplasmic male-sterile line correlates with presence of morphologically modified mitochondria. The putative causes of reduced cell number in the floral meristem and the consequences for floral development are discussed.     

Development of the gametophytes, flower, and floral vasculature in Cochliostema odoratissimum (Commelinaceae)

Flowers of Cochliostema odoratissimum are trimerous with three fertile stamens, three unequal antherless staminodes. and three connate carpels. The fertile stamens are on one side of die flower and united by their filaments, forming a compound structure that curves to the flower's right as the flower opens. The thecae arc longitudinally dehiscent, spirally coiled, and enveloped by pctaloid extensions of the filaments of the two lateral stamens contributing to the three-staminate structure. Anther wall development is of the monocotyledonous type. Tapetal raphides are formed and appear to be widespread in CCommelinaceae. Also known from Philydraceae and. perhaps. Haemodoraccae, tapetal raphides and their taxonomic distribution may be of phylogenetic utilitv. Microsporogcnesis is successive, forming both isobilateral and decussate tetrads. Pollen is shed as single binucleatc grains. Each ovary locule contains ten to twelve hemianatropous, crassinucellar. bitegmic ovules on axile placentae. The micropyle is formed by both the inner integument and one side of the outer integument. Megagamctophyte development is of the Polygonum type. The mature megagamctophyte consists of an egg apparatus, fusion nucleus, and three antipodals. the latter showing signs of degeneration. The salient features of the floral vasculature are the same as in the few other commclinaceous species for winch complete data are available. Relative to the floral vasculature in the other species, differences in the vasculature lie primarily in the presence and origin of lateral carpel bundles and in the number of sepal and ovule traces.     

Origins of radial symmetry identified in an echinoderm during adult development and the inferred axes of ancestral bilateral symmetry

How the radial body plan of echinoderms is related to the bilateral body plan of their deuterostome relatives, the hemichordates and the chordates, has been a long-standing problem. Now, using direct development in a sea urchin, I show that the first radially arranged structures, the five primary podia, form from a dorsal and a ventral hydrocoele at the oral end of the archenteron. There is a bilateral plane of symmetry through the podia, the mouth, the archenteron and the blastopore. This adult bilateral plane is thus homologous with the bilateral plane of bilateral metazoans and a relationship between the radial and bilateral body plans is identified. I conclude that echinoderms retain and use the bilateral patterning genes of the common deuterostome ancestor. Homologies with the early echinoderms of the Cambrian era and between the dorsal hydrocoele, the chordate notochord and the proboscis coelom of hemichordates become evident.     

Effects of Ri TL-DNA from Agrobacterium rhizogenes and the inhibitors of polyamine synthesis on growth, floral development, sexual organogenesis and polyamine metabolism in tobacco

Our recent work associated the phenotypic alterations caused by the Ri TL-DNA (root-inducing, left-hand, transferred DNA) from Agrobacterium rhizogenes with a general decrease in the accumulation of polyamines and their derivatives and showed that irreversible and specific inhibition of putrescine biosynthesis causes a phenotype similar to that attributed to the Ri TL-DNA. In this review we report recent data supporting a correlation between the degree of expression of the transformed phenotype due to the root-inducing, left-hand, transferred DNA and inhibition of polyamine accumulation, strongly suggesting that genes carried by the root-inducing, transferred DNA may act through interference with polyamine production via the ornithine pathway.     

Dwarf and deformed flower 1, encoding an F‐box protein,is critical for vegetative and floral development in rice (Oryza sativa L.)

Recent studies have shown that F‐box proteins constitute a large family in eukaryotes, and play pivotal roles in regulating various developmental processes in plants. However, their functions in monocots are still obscure. In this study, we characterized a recessive mutant dwarf and deformed flower 1‐1 (ddf1‐1) in Oryza sativa (rice). The mutant is abnormal in both vegetative and reproductive development, with significant size reduction in all organs except the spikelet. DDF1 controls organ size by regulating both cell division and cell expansion. In the ddf1‐1 spikelet, the specification of floral organs in whorls 2 and 3 is altered, with most lodicules and stamens being transformed into glume‐like organs and pistil‐like organs, respectively, but the specification of lemma/palea and pistil in whorls 1 and 4 is not affected. DDF1 encodes an F‐box protein anchored in the nucleolus, and is expressed in almost all vegetative and reproductive tissues. Consistent with the mutant floral phenotype, DDF1 positively regulates B‐class genes OsMADS4 and OsMADS16, and negatively regulates pistil specification gene DL. In addition, DDF1 also negatively regulates the Arabidopsis LFY ortholog APO2, implying a functional connection between DDF1 and APO2. Collectively, these results revealed that DDF1, as a newly identified F‐box gene, is a crucial genetic factor with pleiotropic functions for both vegetative growth and floral organ specification in rice. These findings provide additional insights into the molecular mechanism controlling monocot vegetative and reproductive development.     

The relationship between cell division and elongation during development of the nectar-yielding petal spur in Centranthus ruber (Valerianaceae)

Background and Aims Floral spurs are hollow, tubular outgrowths that typically conceal nectar. By their involvement in specialized pollinator interactions, spurs have ecological and evolutionary significance, often leading to speciation. Despite their importance and diversity in shape and size among angiosperm taxa, detailed investigations of the mechanism of spur development have been conducted only recently.Methods Initiation and growth of the nectar-yielding petal spur of Centranthus ruber ‘Snowcloud’ was investigated throughout seven stages, based on bud size and developmental events. The determination of the frequency of cell division, quantified for the first time in spurs, was conducted by confocal microscopy following 4'',6-diamidino-2-phenylindole (DAPI) staining of mitotic figures. Moreover, using scanning electron microscospy of the outer petal spur surface unobstructed by trichomes, morphometry of epidermal cells was determined throughout development in order to understand the ontogeny of this elongate, hollow tube.Key Results Spur growth from the corolla base initially included diffuse cell divisions identified among epidermal cells as the spur progressed through its early stages. However, cell divisions clearly diminished before a petal spur attained 30 % of its final length of 4·5 mm. Thereafter until anthesis, elongation of individual cells was primarily responsible for the spur’s own extension. Consequently, a prolonged period of anisotropy, wherein epidermal cells elongated almost uniformly in all regions along the petal spur’s longitudinal axis, contributed principally to the spur’s mature length.Conclusions This research demonstrates that anisotropic growth of epidermal cells – in the same orientation as spur elongation – chiefly explains petal spur extension in C. ruber. Representing the inaugural investigation of the cellular basis for spur ontogeny within the Euasterids II clade, this study complements the patterns in Aquilegia species (order Ranunculales, Eudicots) and Linaria vulgaris (order Lamiales, Euasterids I), thereby suggesting the existence of a common underlying mechanism for petal spur ontogeny in disparate dicot lineages.     

Cytoskeletal dynamics ofApedinella radians (Pedinellophyceae) III. Post-division development,maintenance of cell symmetry,and the re-establishment of interphase morphology

Summary The dynamics of the cytoskeletal proteins centrin, actin, and tubulin were investigated during post-division development in the radially symmetrical phytoflagellateApedinella radians (Pedinellophyceae). Each daughter cell inherits a triangular arrangement of centrin filamentous bundles that develops, during post-division, into the six-pointed star configuration observed at interphase. This coincides with developmental processes including plaque duplication and migration, chloroplast division and migration, and spine-scale deployment. Centrin filamentous bundles appear to be involved in maintaining radial symmetry throughout the cell cycle and re-establishing interphase morphology. Actin filamentous bundles, prominent at interphase, depolymerize just prior to mitosis and do not reform until late post-division, indicating they are not involved in maintaining cell symmetry during cell division. Although the precise dynamics of microtubular triads and their associated cylindrical caps has not been determined, they may work in concert with centrin filamentous bundles in re-establishing interphase morphology. Three centrin, or centrin-like, components inA. radians appear to coordinate independent architectural events during the cell cycle. The nature of the three centrin components is discussed and compared to the flagellar roots/pericentriolar material of the eukaryotic centrosome.     

Early floral development and androecium organization in the sarracenioid clade (Actinidiaceae,Roridulaceae and Sarraceniaceae) of Ericales

The early floral development of Actinidia (A. arguta, A. callosa, A. chinensis and A. kolomikta; Actinidiaceae), Saurauia (S. montana, S. oldhamii, S. pittieri and S. subspinosa; Actinidiaceae), Roridula gorgonias (Roridulaceae) and Heliamphora nutans (Sarraceniaceae) was studied comparatively using scanning electron microscopy. Late stages of androecium development are additionally presented for Clematoclethra scandens (Actinidiaceae), Darlingtonia californica and Sarracenia leucophylla (Sarraceniaceae). Flowers are typically pentamerous and share a number of developmental features: perianth organs emerge in a clockwise or anticlockwise spiral sequence on the floral apex with relatively long plastochrons between successive organs, resulting in conspicuous size differences among perianth organs in early development; the perianth always consists of two differentiated whorls (unlike earlier interpretations of the perianth in Heliamphora); the androecium is polystemonous in most species and is initiated with leading stamens in alternipetalous positions; successive stamen primordia appear in a lateral succession until a ring‐like structure is formed; and the anthers become inverted shortly before anthesis. Later androecial development differs conspicuously between taxa and further proliferation may be centrifugal, centripetal and/or lateral. For Ericales, unusual features of floral development include: petals initiated in a spiral sequence (but later organized in a whorl) with comparatively long plastochrons between individual petals (except Saurauia); common occurrence of perianth organs in double positions in Actinidiaceae; and anthers that become inverted close to anthesis. The floral development in the sarracenioids is additionally compared with that of other families and clades in Ericales, further emphasizing the highly variable patterns of androecium development in the order.     

The influence of pre-floral growth and development on the pathway of floral development, dry matter distribution and seed yield in oilseed rape (Brassica napus L.)

Patterns of floral development, dry matter distribution and seed yield were examined in winter oilseed rape plants subjected to different pre-floral growth environments. The duration of pre-floral growth and plant size at flower initiation, measured in terms of total mainstem leaf number, were manipulated by varying the temperature between seedling emergence and flower initiation. Exposure of seedlings to low temperatures from cotyledon expansion onwards markedly reduced the duration of pre-floral growth and the number of leaves on the mainstem. The subsequent development pattern of plants was largely dependent on the date of flower initiation and therefore vernalisation requirement. Indeed, the period of growth from flower initiation to maturity, considered on the basis of thermal time, was directly related to the duration of pre-floral growth and mainstem leaf number. The thermal durations of the bud development phase and flowering period in plants exposed to different pre-floral cold treatments but with a common date of flower initiation were similarly linked to these two parameters. Plants exposed to prolonged periods of low temperature treatment from cotyledon expansion onwards initiated fewer mainstem leaves during a relatively short pre-floral growth phase and their yield potential was limited by a reduction in branch and flower numbers. Plants maintained at higher temperatures produced more mainstem leaves during an extended period of pre-floral growth and supported a greater number of branches and flowers. However, this additional yield potential was not realised due to a reduction in seed numbers and mean seed weight. It appeared that seed yield of these plants was limited by increased competition between an excessive number of lower branches and flowers, a problem apparently created by excessive pre-floral growth. Minimal competition for available assimilates between the limited number of branches of plants with a shorter pre-floral growth phase and fewer mainstem leaves, resulted in lower levels of pod abortion, greater seed production and ultimately increased seed yields.