Basic principles of terminal flower formation

Studies of inflorescences of the mutants bractea and terminal flower1 and double mutant bra tfl1 of Arabidopsis thaliana (L.) Heynh. have shown that the presence of a developed leaf in the node preceding the terminal flower is a necessary condition for the formation of the terminal flower perianth. This means that perianth cannot develop in an abracteose inflorescence of terminal flower. The second necessary condition for the terminal flower formation is a sufficient level of expression of the genes responsible for floral morphogenesis. Combination of these two conditions suffices for the development of a terminal flower with perianth. Since the general principles of organization are common for the majority of Angiosperms, it can be stated that if the abracteose inflorescence is terminated by a flower with perianth, this is a consequence of displacement of the lateral flower into the terminal position.__________Translated from Ontogenez, Vol. 36, No. 2, 2005, pp. 90–95.Original Russian Text Copyright © 2005 by Penin, Choob, Ezhova.     

外源激素在诱导风信子花被外植体不同部位细胞再生花芽中的作用

外源激素诱导风信子（Ｈｙａｃｉｎｔｈｕｓ ｏｒｉｅｎｔａｌｉｓＬ．）同一发育时期花被外植体不同部位细胞再生花芽的实验表明∶１． 诱导花被外植体细胞再生花芽,外源激素是必需的;２． 仅有细胞分裂素就可以诱导花芽再生,生长素并不是必需的;３． 花被外植体上的不同部位的细胞再生花芽时,需要不同浓度的外源激素． 单独加６－ＢＡＰ或玉米素２ ｍ ｇ／Ｌ可以诱导花被下部的细胞再生花芽;６－ＢＡＰ或玉米素２ ｍ ｇ／Ｌ和２,４－Ｄ ０．１ ｍ ｇ／Ｌ的组合有利于花被中部的细胞再生花芽;６－ＢＡＰ或玉米素２ ｍ ｇ／Ｌ和２,４－Ｄ １．０ ｍ ｇ／Ｌ的组合能促进花被上部的细胞分化花芽     

Comparative ontogeny of the cyathium in Euphorbia (Euphorbiaceae) and its allies: exploring the organ-flower-inflorescence boundary

We present a detailed comparative ontogenetic analysis of pseudanthia of representatives of all three subtribes of Euphorbieae (Euphorbiinae, Neoguillauminiinae, Anthosteminae) in order to clarify their homologies and interpretation. The cyathium of Euphorbia and its allies (subtribe Euphorbiinae) closely resembles a bisexual flower but is traditionally interpreted as an inflorescence bearing clusters of highly reduced male flowers surrounding a single terminal female flower. Previously unreported characters are (1) male flowers formed one above the other in the male inflorescences of some Euphorbiinae, (2) late-developing perianthlike structures in some male flowers of Neoguillauminia cleopatra, (3) evidence for a bracteate origin of the female perianth in Anthosteminae and Neoguillauminiinae, and (4) spatiotemporally independent formation of abscission zone and perianth. Indistinct boundaries between inflorescence, flower, and floral organs demonstrate that defining the cyathium neither as an inflorescence nor as a flower is entirely satisfactory and indicate a "hybrid" flower/inflorescence nature of the cyathium. Based on our current knowledge and the existing phylogenetic context, it is most parsimonious to suggest that the cyathium evolved from a determinate thyrse with a terminal female flower surrounded by dichasial male partial inflorescences. We speculate that the cyathium was formed because of strong condensation and possible overlap between expression zones of regulatory genes.     

Role of Exogenous Hormones in Inducing Cells in different Positions of Perianth Explants to Regenerate Flower Buds in Hyacinthus orientalis

Role of the exogenous hormone in inducing different position cells of perianth explants of hyacinth to regenerate flower buds was studied. Experiments showed that (1) Exogenous hormones are necessary for inducing cells of the perianth explant to regenerate the flower buds; (2) Only cytokinine alone could induce the regeneration of the flower buds, the auxin was not necessary; (3) Exogenous hormones in different concentrations could induce cells in the different parts of the perianth explants to differentiate the flower buds: 6-BAP or zeatin 2 mg/L alone could induce cells located at the lower part of the perianth to differentiate flower buds. Combination of 6-BAP or zeatin 2 mg/L and 2, 4-D 0.1 mg/L was advantageous to cells located middle part of the perianth to regenerate the flower buds. Combination of 6-BAP or zeatin 2 mg/L and 2, 4-D 1.0 mg/L could promote cells located at the upper part of the perianth to differentiate flower buds.     

Intra‐individual variation of flowers in Gunnera subgenus Panke (Gunneraceae) and proposed apomorphies for Gunnerales

A study of inflorescence and flower development in 12 species from four of the six subgenera of Gunnera (Gunneraceae) was carried out. In the species of subgenus Panke, initiation of floral apices along the partial inflorescences is acropetal but ends up in the late formation of a terminal flower, forming a cyme at maturity. The terminal flower is the largest and the most complete in terms of merosity and number of whorls and thus it is the most diagnostic in terms of species‐level taxonomy. The lateral flowers undergo a basipetal gradient of organ reduction along the inflorescence, ranging from bisexual flowers (towards the distal region) to functionally (i.e. with staminodia) and structurally female flowers (towards the proximal region). Our results show that the terminal structure in Gunnera is a flower rather than a pseudanthium. The terminal flower is disymmetric, dimerous and bisexual, representing the common bauplan for Gunnera flowers. It has a differentiated perianth with two sepals and two alternate petals, the latter opposite the stamens and carpels. Comparisons with other members of the core eudicots with labile floral construction are addressed. We propose vegetative and floral putative synapomorphies for the sister‐group relationship between Gunneraceae and Myrothamnaceae. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 262–283.     

粤北南方红豆杉的群落类型及物候与繁殖生物学特性

对粤北地区的南方红豆杉（Taxus maikrei）植物进行物候学观察,以有植物群落调查,并着重研究其种子萌发和插插繁殖特性,结果表明,在粤北地区,南方红豆杉主要分布于南亚热带中亚热带典型的常绿阔叶林中;种群个体顶芽展梢在4月中旬开始,5月中旬雄球花现蕾,花期7月底至11月下旬,8月下旬雌球花现蕾,花期10月下旬至次年1月底,果期在次年10月上旬,10月下旬顶芽形成;在自然条件下种子需2－3年才能萌发;在山地条件下对种子进行湿沙藏,再播入沙床中,萌发率达到95．2％,但周期较长（1年半）;在实验条件下对种子采用为温层积（先25℃36周,再转入5℃12周,共48周,共48周,即336d）,取出培养萌发率最高,达82．2％;扦插繁殖方面,以成熟一年生枝为插穗,用生根粉ABT1浓度在100mg.L^-1时处理效果较好,扦插时间在10－11月份最适合,生根率达95％。     

PERIANTH DEVELOPMENT OF POTAMOGETON RICHARDSONII

Interpretation of the Potamogeton flower is complicated by the attachment of the “perianth segment” to the stamen connective. Developmental studies show that the perianth segments are not outgrowths of the stamen connectives. They are initiated on the floral apex acropetally before the (superposed) primordia of the stamens. After the inception of the stamen primordia, growth occurs in the regions between the primordia of each perianth segment and stamen. Thereby the bases of the developing perianth segment and stamen become united, and in the adult flower eventually the perianth segment is inserted on the connective of the stamen. The primordium of the perianth segment develops from the 2 outer layers (tunica) of the floral apex, in contrast to the stamen primordium which originates from the 3 outer layers. The vascular bundles for each perianth segment–stamen region develop acropetally from 1 common bundle which bifurcates into 1 bundle for the perianth segment and 1 for the stamen. The bundle leading into the perianth segment branches in a more or less dichotomous manner. The veins form none or only 1 or 2 anastomoses at the base of the lamina, whereas the vein endings remain free. The interpretation of the perianth segments is discussed in terms of the classical and the gonophyll theory. Since both theories rest on an ambiguous methodological basis, interpretation is postponed until a new approach to comparative morphology has been worked out and until the floral development of other Helobiales has been studied.     

Type specification and spatial pattern formation of floral organs: A dynamic development model

A mathematical model simulating spatial pattern formation (positioning) of floral organs is proposed. Computer experiment with this model demonstrated the following sequence of spatial pattern formation in a typical cruciferous flower: medial sepals, carpels, lateral sepals, long stamens, petals, and short stamens. The positioning was acropetal for the perianth organs and basipetal for the stamens and carpels. Organ type specification and positioning proceed non-simultaneously in different floral parts and organ type specification goes ahead of organ spatial pattern formation. Computer simulation of flower development in several mutants demonstrated that the AG and AP2 genes determine both organ type specification and formation of the zones for future organ development. The function of the AG gene is to determine the basipetal patterning zones for the development of the reproductive organs, while the AP2 gene maintains proliferative activity of the meristem establishing the acropetal patterning zone for the development of the perianth organs.     

Induction of Continuous Tepal Differentiation from in Vitro Regenerated Flower Buds of Hyacinthus orientalis

Continuous differentiation of tepals was successively induced from regenerated flower buds in Hyacinthus orientalis L. cv. White Pearl by controlling the exogenous hormones and explant ages. In 250 days of subculture, each flower bud differentiated an average of more than 70 tepals, with a maximum of over 140 tepals. Studies on the morphogenesis and characteristics of growth and development of the flower buds indicate that the first whorled organ of the flower bud was perianth which consisted of perianth tube and tepals grown at the top of the perianth tube, which is the same as the flower bud of the wild type in H. orentalis. The second and third whorls of the flower bud, which should be stamen and pistil in the wild type, but remained as the tepals in the regenerated flower bud. Growth of the regenerated flower bud was faster in the first several months of culture, then slowed down gradually with time. After 150 days in culture the flower bud growth and organ differentiation became very slow. Other than the tepal differentiation the regenerated flower buds also differentiated at random positions some small flower buds that also differentiated the tepals only. Histological observation revealed that the origin of the regenerated flower buds was jointly participated by some cells in the epidermal and subepidermal layers at the inner surface of the perianth explant, and the inner small flower buds were originated from the meristem which was formed by the transformation of the parenchyma at the base of the very young tepal. The authors also compared and discussed the similarities and differences of the phenotypes between the regenerated flower bud in Hyacinthus and agamous flower in Arabidopsis, from which, they have hypothesized on the role of the hormones in the promotion and termination of the gene expressions by an order of development in plant.     

紫花含笑花被呈色过程中色素含量与超微结构的变化特征

为了探究色素含量以及细胞结构在紫花含笑花被呈色过程中的作用机理,该研究以绿色和紫色花被为材料,测定其花被色素含量,运用逐步回归方程分析花被呈色与色素含量的关系,采用石蜡切片及超薄切片技术观察花被细胞超显微结构变化.结果表明:(1)在紫花含笑花被呈色过程中,紫色花被表面明度L*值降低,a*值上升,b*值降低;花被花青素苷...     

FLORAL DEVELOPMENT IN MYRISTICA (MYRISTICACEAE)

Myristica fragrans and M. malabarica are dioecious. Both staminate and pistillate plants produce axillary flowering structures. Each pistillate flower is solitary, borne terminally on a short, second-order shoot that bears a pair of ephemeral bracts. Each staminate inflorescence similarly produces a terminal flower and, usually, a third-order, racemose axis in the axil of each pair of bracts. Each flower on these indeterminate axes is in the axil of a bract. On the abaxial side immediately below the perianth, each flower has a bracteole, which is produced by the floral apex. Three tepal primordia are initiated on the margins of the floral apex in an acyclic pattern. Subsequent intercalary growth produces a perianth tube. Alternate with the tepals, three anther primordia arise on the margins of a broadened floral apex in an acyclic or helical pattern. Usually two more anther primordia arise adjacent to each of the first three primordia, producing a total of nine primordia. At this stage the floral apex begins to lose its meristematic appearance, but the residuum persists. Intercalary growth below the floral apex produces a columnar receptacle. The anther primordia remain adnate to the receptacle and grow longitudinally as the receptacle elongates. Each primordium develops into an anther with two pairs of septate, elongate microsporangia. In pistillate flowers, a carpel primordium encircles the floral apex eventually producing an ascidiate carpel with a cleft on the oblique apex and upper adaxial wall. The floral ontogeny supports the morphological interpretation of myristicaceous flowers as trimerous with either four-sporangiate anthers or monocarpellate pistils.     

Does Ethylene Treatment Mimic the Effects of Pollination on Floral Lifespan and Attractiveness?

In some species pollination may result in rapid changes in perianth colour and form (petal senescence and abscission, flower closure), rendering the flowers less attractive to pollinators. It has been suggested that this effect is mediated by ethylene. Flowers from about 200 species and 50 families were exposed to ethylene (3 ppm for 24 h at 20 degrees C). The effects on petal senescence and abscission have been described previously. Flower closure and perianth colour changes were generally ethylene-sensitive, but responses showed no consistency within families. Several flowers known to respond to pollination by rapid cessation of attractiveness were also exposed to ethylene: this produced the same effect as pollination, both on flower colour and form. Species that respond to pollination by changing flower form or colour were found exclusively in families in which the species are generally ethylene-sensitive (with regard to changes in perianth form and colour). However, several families are generally ethylene-sensitive but contain no species reported to respond to pollination.     

Inflorescence and floral ontogeny in Crocus sativus L. (Iridaceae)

Inflorescence and floral ontogeny of the perennial, herbaceous crop Crocus sativus L. were studied using epi-illumination light microscopy. After production of leaves with helical arrangement a determinate inflorescence forms which becomes completely transformed into a single terminal flower. In some cases, bifurcation of the inflorescence meristem yields two or three floral meristems. The order of floral organs initiation is outer tepals – stamens – inner tepals – carpels. Stamens and outer tepals are produced from the lateral bifurcation of three common stamen-tepal primordia. Within each whorl, organs start developing unidirectionally from the adaxial side, except for the stamens which begin to grow from the abaxial side. Specialized features during organ development include interprimordial growth between tepals forming a perianth tube, fusion at the base of stamen filaments, and formation of an inferior ovary with unfused styles.     

Floral Morphology of Populus lasiocarpa Oliv and Its Phylogenetic Position in Populus

The results of this study, which involved macroscopic, microscopic and SEM investigation of flowers of Populus lasiocarpa Oliv., may be stated as follows: 1. Female, male and bisexual flowers possess well developed perianth greenish in color with obvious veins and glandular teeth. The branched vascular bundles and stomata apparatus are observed in the transection of the perianth. It suggests that the perianth is homologous with the leaf. 2. Each pistil of female and bisexual flowere possesses 3(–5) separate styles. The ovary is I-celled with 16–34 ovules. Each male flower has 41–110 stamens. The connective of stamen is protrusive. Presence of bisexual, polygamous flower and monoecism is not rare. 3. In comparison with several other species of Populus, P. lasiocarpa is supposed to be the most ancient and primitive species of the Populus. The central and south-west China perhaps is one of distributional center of Populus.     

Isolation of the tomato AGAMOUS gene TAG1 and analysis of its homeotic role in transgenic plants.

To understand the details of the homeotic systems that govern flower development in tomato and to establish the ground rules for the judicious manipulation of this floral system, we have isolated the tomato AGAMOUS gene, designated TAG1, and examined its developmental role in antisense and sense transgenic plants. The AGAMOUS gene of Arabidopsis is necessary for the proper development of stamens and carpels and the prevention of indeterminate growth of the floral meristem. Early in flower development, TAG1 RNA accumulates uniformly in the cells fated to differentiate into stamens and carpels and later becomes restricted to specific cell types within these organs. Transgenic plants that express TAG1 antisense RNA display homeotic conversion of third whorl stamens into petaloid organs and the replacement of fourth whorl carpels with pseudocarpels bearing indeterminate floral meristems with nested perianth flowers. A complementary phenotype was observed in transgenic plants expressing the TAG1 sense RNA in that first whorl sepals were converted into mature pericarpic leaves and sterile stamens replaced the second whorl petals.     

Male and female flowers of the dioecious plant sorrel show different patterns of MADS box gene expression.

Male and female flowers of the dioecious plant sorrel (Rumex acetosa) each produce three whorls of developed floral organs: two similar whorls of three perianth segments and either six stamens (in the male) or a gynoecium consisting of a fertile carpel and two sterile carpels (in the female). In the developing male flower, there is no significant proliferation of cells in the center of the flower, in the position normally occupied by the carpels of a hermaphrodite plant. In the female flower, small stamen primordia are formed. To determine whether the organ differences are associated with differences in the expression of organ identity genes, cDNA clones representing the putative homologs of B and C function MADS box genes were isolated and used in an in situ hybridization analysis. The expression of RAD1 and RAD2 (two different DEFICIENS homologs) in males and females was confined to the stamen whorl; the lack of expression in the second, inner perianth whorl correlated with the sepaloid nature of the inner whorl of perianth segments. Expression of RAP1 (a PLENA homolog) occurred in the carpel and stamen whorls in very young flower primordia from both males and females. However, as soon as the inappropriate set of organs ceased to develop, RAP1 expression became undetectable in those organs. The absence of expression of RAP1 may be the cause of the arrest in organ development or may be a consequence.     

Visual discrimination between two sexually deceptive <Emphasis Type="Italic">Ophrys</Emphasis> species by a bee pollinator

Almost all species of the orchid genus Ophrys are pollinated by sexual deception. The orchids mimic the sex pheromone of receptive female insects, mainly hymenopterans, in order to attract males seeking to copulate. Most Ophrys species have achromatic flowers, but some exhibit a coloured perianth and a bright, conspicuous labellum pattern. We recently showed that the pink perianth of Ophrys heldreichii flowers increases detectability by its pollinator, males of the long-horned bee Eucera berlandi. Here we tested the hypothesis that the bright, complex labellum pattern mimics the female of the pollinator to increase attractiveness toward males. In a dual-choice test we offered E. berlandi males an O. heldreichii flower and a flower from O. dictynnae, which also exhibits a pinkish perianth but no conspicuous labellum pattern. Both flowers were housed in UV-transmitting acrylic glass boxes to exclude olfactory signals. Males significantly preferred O. heldreichii to O. dictynnae flowers. In a second experiment, we replaced the perianth of both flowers with identical artificial perianths made from pink card, so that only the labellum differed between the two flower stimuli. Males then chose between both stimuli at random, suggesting that the presence of a labellum pattern does not affect their choice. Spectral measurements revealed higher colour contrast with the background of the perianth of O. heldreichii compared to O. dictynnae, but no difference in green receptor-specific contrast or brightness. Our results show that male choice is guided by the chromatic contrast of the perianth during the initial flower approach but is not affected by the presence of a labellum pattern. Instead, we hypothesise that the labellum pattern is involved in aversive learning during post-copulatory behaviour and used by the orchid as a strategy to increase outcrossing.     

Inflorescence and floral development in <Emphasis Type="Italic">Trochodendron aralioides</Emphasis> (Trochodendraceae)

In the early development of Trochodendron aralioides (Trochodendraceae) inflorescences lateral flowers are initiated after the appearance of the floral pherophylls (subtending bracts). The terminal flower is preceded by metaxyphylls and is initiated earlier than the uppermost lateral flowers of the botryoid inflorescence. Small scales (interpreted as rudimentary perianth organs) precede the stamens. These scales are more distinct in the terminal flower than in the lateral flowers. In the radially symmetrical terminal flower, small scales (or metaxyphylls) and stamens are initiated in a spiral during early development. At anthesis, stamen phyllotaxis appears irregular or approximately whorled as a result of the rapid elongation and irregular slight curvature of the stamen filaments which distorts the originally regular pattern. Finally, the numerous carpels arise simultaneously in a single whorl. It takes about 9 months for flowers to develop and the 2-year reproductive cycle of T. aralioides is typical of many trees. The floral development of T. aralioides is compared with that of other basal eudicots. The bottle-shaped, unicellular stigmatic papillae and long, decurrent stigma of basally united carpels are similar to those of the Buxales¸ suggesting a close relationship.     

Chaotic Floral Phyllotaxis and Reduced Perianth in Achlys (Berberidaceae)

Among the 16 genera of the Berberidaceae Achlys is the only one with a reduced perianth, an irregular floral phyllotaxis, and variable stamen number. Early floral stages show an unstable (chaotic) arrangement of the organ primordia. Only the single carpel of the gynoecium has a more fixed position in that the placenta is formed in the adaxial half of the flower. The irregularities in the androecium may be caused by the lack of influence of a perianth on floral symmetry. On the other hand, the regular orientation of the carpel is perhaps due to the early polarity of the flower, whereby the abaxial half of the flower is larger (with further developed stamen primordia) at the time when carpel polarity is established.     

A Mutation in the Arabidopsis TFL1 Gene Affects Inflorescence Meristem Development

We present the initial phenotypic characterization of an Arabidopsis mutation, terminal flower 1-1 (tfl1-1), that identifies a new genetic locus, TFL1. The tfl1-1 mutation causes early flowering and limits the development of the normally indeterminate inflorescence by promoting the formation of a terminal floral meristem. Inflorescence development in mutant plants often terminates with a compound floral structure consisting of the terminal flower and one or two subtending lateral flowers. The distal-most flowers frequently contain chimeric floral organs. Light microscopic examination shows no structural aberrations in the vegetative meristem or in the inflorescence meristem before the formation of floral buttresses. The wild-type appearance of lateral flowers and observations of double mutant combinations of tfl1-1 with the floral morphogenesis mutations apetala 1-1 (ap1-1), ap2-1, and agamous (ag) suggest that the tfl1-1 mutation does not affect normal floral meristems. Secondary flower formation usually associated with the ap1-1 mutation is suppressed in the terminal flower, but not in the lateral flowers, of tfl1-1 ap1-1 double mutants. Our results suggest that tfl1-1 perturbs the establishment and maintenance of the inflorescence meristem. The mutation lies on the top arm of chromosome 5 approximately 2.8 centimorgans from the restriction fragment length polymorphism marker 217.