Theligonum cynocrambe: Developmental morphology of a peculiar rubiaceous herb

The annual Mediterranean herbTheligonum cynocrambe shows a peculiar combination of morphological characters, e.g., switch from decussate to spiral phyllotaxis with 90–100° divergence, combined with a change from interpetiolar to lateral stipules, anemophily, lack of calyx, flowers often dimerous to trimerous, corolla fused in both male and female flowers, male flowers extra-axillary, with 2–19 stamens per flower, female flowers axillary, with inferior uniovulate ovary, basilateral style and perianth, nut-like fruits with elaiosome. In male flowers the androecium emerges as an (uneven) elliptical rim with a central depression. This common girdling primordium is divided up into several stamen primordia. In male flowers with low stamen number the stamen primordia may occupy the corners alternating with the corolla lobes. There are no epipetalous androecial primordia that secondarily divide into stamens. Male flowers occasionally show a hemispherical base that may be interpreted as remnant of the inferior ovary. In female flowers a ring primordium grows into a tube on which the petal lobes arise. The perianth and style become displaced adaxially by uneven growth of the inferior ovary. The ovary is basically bilocular. The lower region of the ovary is provided with a septum that is overtopped and hidden by the single curved ovule.Theligonum is referred to theRubiaceae-Rubioideae, with theAnthospermeae andPaederieae as most closely related tribes.     

Development of male and female flower in Asparagus officinalis. Search for point of transition from hermaphroditic to unisexual developmental pathway

Asparagus officinalis is a dioecious plant. The flowers start to develop as hermaphrodites and later become unisexual. In female flowers the stamens degenerate, while in male flowers the ovary stops growing without degenerating. We have examined young asparagus flowers using SEM and optical microscopy in order to determine the exact moment of transition from hermaphroditic to unisexual development. We defined 13 stages of development, starting from flower primordia up to completely mature flowers and labelled them with numbers from -6 to 7. The first five stages are fully hermaphroditic: a difference between sexes becomes visible at stage — 1 when the style begins to develop in female flowers. Degeneration of stamens in female flowers starts somewhat later. At the stage of transition, some differences between sexes also appear in the bidimensional polypeptide pattern of flowers. RNase activity shows a distinct peak at this stage (in female flowers only), probably related to stamen degeneration.     

河南省五加科1新记录属——梁王茶属

报道了河南省五加科(Araliaceae)1个分布新记录属及其属下1新记录种,梁王茶属(Metapanax J. Wen & Frodin),异叶梁王茶［Metapanax davidii (Franchet) J. Wen & Frodin］。该种为常绿灌木,叶为单叶或掌状分裂,花聚生为伞形花序,再组成顶生的圆锥花序,苞片早落,花梗具关节,子房2室,核果,扁球形,花期7～8月,果期9～11月。讨论了该种叶、花、果的形态学变异特征和其区系地理学意义。该新记录的发现对梁王茶属的系统分类、迁移演化以及药用植物资源开发具有重要的意义。     

Two-dimensional patterns of soluble proteins including three hydrolytic enzymes of mature pollen of tristylous <Emphasis Type="Italic">Lythrum salicaria</Emphasis>

Lythrum salicaria, now a widespread invasive species, exhibits tristyly, a form of heteromorphic selfincompatibility. In tristyly, each plant exhibits one (and only one) of three morphologically different floral forms. Moreover, each flower produces two types of stamens, and these two exhibit different incompatibility reactions. Differences between stamens of a single flower must be the result of epigenetic phenomena and for that reason, we performed two-dimensional gel electrophoresis (2-DE) to analyze fractions of soluble proteins derived from the pollen coat and protoplast including three hydrolytic enzymes from the six different stamen types (two from each of three floral forms). There were significant differences in the 2-D protein profiles both between pollen from the same flower and between the same type of pollen from two different flowers, in the pollen coat as well as in the protoplast extracts. In five of the six samples of pollen fractions, characteristic peptides were found. Quantitative differences between pollen from the same flower were observed in case of esterases. Furthermore, analysis of proteases and acid phosphatases revealed also qualitative differences between these enzymes in pollen from the same flower.     

商陆属2种植物花器官发生过程观察

该研究采用扫描电镜观察红蕊商陆(Phytolacca esculenta)和浙江商陆(Phytolacca zhejiangensis)的花器官发生过程,以明确商陆属植物花的基数,以及雄蕊和雌蕊是否具有叶性器官发生的特点,阐明商陆属植物花发生的模式。观察结果显示:(1)红蕊商陆和浙江商陆在花原基发生后,小苞片以2/5圆周相继发生,花被片的发生紧接小苞片的发生进行,花被与小苞片的发生均有顺时针和逆时针方向,且二者的发生方向始终一致。(2)花被发生结束后,雄蕊在花顶端分生组织的环状分生组织上发生,没有明显的发生顺序,近似同时发生;2轮雄蕊时内轮雄蕊先发生;外轮雄蕊有少数有时偶然与花被互生,但因外轮雄蕊数多于花被数,雄蕊与花被常不互生,也没有规律性。(3)红蕊商陆和浙江商陆的心皮都在雄蕊发生后,紧接着开始发生,且雌蕊与雄蕊(或内轮雄蕊)互生发生;心皮没有发生的先后次序,且每个心皮在基部连成一个整体形成雌蕊基部并发育成为子房。(4)红蕊商陆和浙江商陆的花基数为5,雄蕊和雌蕊的发生及数目不符合5基数的特点。研究认为,红蕊商陆和浙江商陆为5基数花,该研究结果不支持商陆属植物为3基数花的发生模式。     

Relationships between floral organization, architecture, and pollination mode inDillenia (Dilleniaceae)

The flowers ofDillenia are highly elaborate pollen-flowers adapted to buzzpollination byXylocopa bees. Two major forms of floral architecture (revolver flowers and roundabout flowers) are associated with two different pollination modes. In the first (e.g.,D. suffruticosa), the pollination organs are connivent to a cone; the pollinator grasps the entire cone with its legs and buzzes it; it revolves around its axis and repeats the buzzing in different positions. In the second (e.g.,D. alata, D. philippinensis), the stylar branches are spreading and the stamens are arranged in two sets of two different forms and colourations. The inner set has fewer and longer stamens that are cryptic pollination stamens; those of the outer set are shorter but optically conspicuous feeding stamens. The pollinator squeezes itself under the stylar branches and handles only the outer set by grasping part of the set at a time; it moves tangentially around the flower with several buzzing-stops; when buzzing pollen is sprayed onto its side and back from the inner stamen set. Centrifugal polyandrous androecia are a constitutive feature of flowers inDilleniaceae. InDillenia the centrifugal initiation of stamens proceeds for an unusually long time and is still not finished when the gynoecium is completely closed (in contrast toTetracera). The differentiation of heteranthery seems to be functionally correlated with the extended centrifugal inception. The latest formed stamens are small and sterile in many species. Generic features ofDillenia flowers can be understood from the roundabout architecture: big size, increased number of carpels, syncarpy forming a firm pedestal and spreading firm stylar branches with small, concave stigmas at the end, stamens with short, stout filaments and much elongated poricidal anthers, heteranthery, recurved stamens of the inner set.Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

洋葱花器数目变异研究

以洋葱JQS-1、MST-140、Red beauty 3个常规栽培品种的鳞茎为试材,观察统计洋葱花器数目的变化,探讨花器数目的变异与植物花器多样性的关系,为植物花器发育模式以及植物分类和系统发育研究提供依据。结果表明:(1)洋葱花序中正常的洋葱小花含有6枚雄蕊,一些异常小花雄蕊的数目减少到5枚或增加到7~11枚;正常花药中的花粉粒数量大、形状规则、分散均匀,而异常花药中的花粉粒形状不规则。(2)雌蕊数目也发生变异,子房由正常的3室,变成了2室、4室或6室;花柱从正常1枚,增加到2枚或3枚。(3)正常小花含有6片花被片,异常小花的花被片数目从5片到10片。(4)花被片与雄蕊数目的变异有同步增减和非同步增加2种类型。     

Pollination biology of the saprophytic speciesPetrosavia sakuraii (Makino) van Steenis in Central Japan

Pollination biology of the saprophytic speciesPetrosavia sakuraii was investigated in Central Japan. The flower opens at any time of the day, and the anthers of outer stamens dehisce after about one day and those of the inner stamens after two days. The stigmata on the semicarpous (nearly apocarpous) pistil, which are already receptive when the flower opens, are situated in the center of the flower and move gradually outwards to touch the anthers of outer stamens about five days after anthesis. The breeding experiments show that the stigmata are receptive at least for five days, and many seeds are produced through autonomous self-pollination. That means the nearly apocarpous pistil, which is in a primitive condition, is adapted to the autonomous self-pollination. The selfing rate including insect-mediated self-pollination is very high, and this sexual-reproductive system seems favorable to the saprophytic plant which is probably severely limited in its resources. However, cross-pollination also may be performed by the small bees of Lasioglossum and some other insects, even though they do not so frequently visit the flowers and the out-crossing rate is low.     

Expression of ASK1-like genes in arrested stamens of female Silene latifolia plants

Silene latifolia is a dioecious plant in which sex is determined by heteromorphic sex chromosomes. In female plants, stamen development is arrested before microspore mother cells are formed. In this study, we isolated four cDNAs (SlSKP1-1 to 4) encoding ASK1-like protein as expression markers to reveal when expression levels are reduced in arrested stamens of female flowers. Expression patterns of the SlSKP1 genes were analyzed by in-situ hybridization. We use the flower development classification of Grant et al. (in Plant J 6:471–480, 1994). SlSKP1 genes were highly expressed in primary parietal cells and primary sporogenous cells that develop into microspore mother cells in male flowers. Expression levels started to be reduced in the external stamens of the female flowers when stamen development was arrested at stage 7. Although microspore mother cells could not be developed in female flowers and SlSKP1 expression may be unnecessary in arrested stamens, SlSKP1 genes were still expressed in sporogenous cells of degenerated stamens at stage 8. Parietal cells stopped differentiating earlier than sporogenous cells in arrested stamens. These results suggest that not all types of cell are arrested simultaneously at a particular stage of stamen development during stamen suppression of S. latifolia.     

Gender expression and inflorescence structure of Pappea capensis Eckl. and Zeyh. (Sapindaceae)

Gender and the structure of the inflorescence and flowers of Pappea capensis (Sapindaceae) are investigated in a locality around Pretoria (22-27°S and 25-32°E). The trees flower over a long period (December to April) and are basically monoecious, starting with male flowers followed by female flowers towards the end of the flowering period, although some trees may be predominantly male and some predominantly female. The inflorescence is a reduced thyrse with small flowers. Male flowers have five ephemeral petals, eight stamens and a rudimental pistil. Female flowers comprise a 3-lobed ovary, a single style and stigma and eight staminodes.     

Gender variation in Actinidia deliciosa,the kiwifruit

Summary Flower and fruit characters were measured in ten female, five male and five fruiting male selections of A. deliciosa var deliciosa (A. Chev) Liang and Ferguson. Flowers from female vines had functional pistils, which contained many ovules. Stamens appeared to be fully developed but produced only empty pollen grains. Flowers from male vines had functional stamens that produced high percentages of pollen grains with stainable cytoplasmic contents. Pistils did not contain ovules and were generally small with vestigial styles. Fruiting male vines had both staminate and bisexual flowers. Staminate flowers were similar to those found on strictly male vines. Bisexual flowers produced ovules and stainable pollen. Pistils were smaller than in pistillate flowers. Although the three flower sexes differed in style length, ovary dimensions and ovules per carpel, staminate and bisexual flowers were similar in number of flowers per inflorescence, stamen filament length, pollen stainability, inflorescence rachis length and carpel number, and differed from pistillate flowers in these characters. The three flower sexes had similar sepal and petal numbers. The fruit of fruiting males were considerably smaller than those of females. Low ovule number appears to be the major factor limiting fruit size in the fruiting males studied. Prospects for developing hermaphroditic kiwifruit cultivars through breeding are discussed.     

桂味荔枝花器官的发生和发育过程研究

利用SV11立体显微镜和JSM-6360LV型扫描电镜观察‘桂味’荔枝花器官的发生和发育过程。结果表明:花序原基最先发生,然后形成数个大小不等的单花原基;4个萼片原基的发生不同步,其中一侧对位先发生;6～10枚雄蕊原基以轮状方式几乎同时发生;心皮原基最后发生,2～3枚(稀4枚)心皮原基同时出现,随后进行侧向生长,逐渐合拢形成子房。雌花中,花柱、柱头分化明显,雄蕊退化。雄花中,花丝细长,花药饱满,雌蕊退化或发育不完全。两性花中,雌雄蕊发育完全。花粉粒近球形,具3孔沟,表面为条纹状纹饰。     

Leafless autumnal-flowering geophytes in the Mediterranean region — phytogeographical,ecological and evolutionary aspects

Numerous Mediterranean geophytes are hysteranthous, i.e. they flower in a leafless stage during autumn. On the basis of their life cycles and other features two types can be recognized: In the Crocus type flowers with subterranean ovary develop at the start of the new reproductive cycle from an annual storage organ; seed dispersal and germination are delayed to the spring and the next autumn. In the Urginea type, flowers with supraterranean ovary originate from a perennial storage organ at the end of the life cycle; seed dispersal and germination follow flowering immediately. Similar rhythms exist in other seasonal climates. Field observations and some experiments demonstrate flower stimulation by temperature decrease or increase in different geophytes, and relationships between water supply and synanthous versus hysteranthous leaf development. The origin, ecological importance, and horticultural implication of the hysteranthous syndrome is discussed.     

Carpeloidy in flower evolution and diversification: a comparative study in Carica papaya and Arabidopsis thaliana

Background and Aims

Bisexual flowers of Carica papaya range from highly regular flowers to morphs with various fusions of stamens to the ovary. Arabidopsis thaliana sup1 mutants have carpels replaced by chimeric carpel–stamen structures. Comparative analysis of stamen to carpel conversions in the two different plant systems was used to understand the stage and origin of carpeloidy when derived from stamen tissues, and consequently to understand how carpeloidy contributes to innovations in flower evolution.

Methods

Floral development of bisexual flowers of Carica was studied by scanning electron microscopy and was compared with teratological sup mutants of A. thaliana.

Key Results

In Carica development of bisexual flowers was similar to wild (unisexual) forms up to locule initiation. Feminization ranges from fusion of stamen tissue to the gynoecium to complete carpeloidy of antepetalous stamens. In A. thaliana, partial stamen feminization occurs exclusively at the flower apex, with normal stamens forming at the periphery. Such transformations take place relatively late in development, indicating strong developmental plasticity of most stamen tissues. These results are compared with evo-devo theories on flower bisexuality, as derived from unisexual ancestors. The Arabidopsis data highlight possible early evolutionary events in the acquisition of bisexuality by a patchy transformation of stamen parts into female parts linked to a flower axis-position effect. The Carica results highlight tissue-fusion mechanisms in angiosperms leading to carpeloidy once bisexual flowers have evolved.

Conclusions

We show two different developmental routes leading to stamen to carpel conversions by late re-specification. The process may be a fundamental aspect of flower development that is hidden in most instances by developmental homeostasis.     

Stamen movements in flowers ofOpuntia (Cactaceae) favour oligolectic pollinators

Opuntia brunneogemmia andO. viridirubra occur sympatrically in the Serra do Sudeste, Rio Grande do Sul, Brazil. Their flowers have 450–600 thigmonastic stamens and provide large amounts of pollen and nectar for bees. Bees of 41 species were registered at the flowers ofO. brunneogemmia and 30 at the flowers ofO. viridirubra. Females of three oligolectic species are the only effective pollinators:Ptilothrix fructifera (Anthophoridae),Lithurgus rufiventris (Megachilidae), andCephalocolletes rugata (Colletidae). During their visits inOpuntia-flowers, bees touch the filaments and stimulate the movement of the stamens to the centre of the flower. At the end of this movement, the anthers are densely packed around the style. As a consequence the pollen is presented in an easily accessible upper layer of anthers and various, nearly inaccessible lower layers. The lower layers contain about 80% of the pollen reward. Only females of the three oligolectic pollinators exploit the pollen from the lower layers and reach the nectar furrow. Therefore, through their stamen movements,Opuntia flowers hide most of their pollen from flower visitors but favour effectively pollinating, oligolectic bees.     

Floral organogenesis in Tetracentron sinense (Trochodendraceae) and its systematic significance

In Tetracentron sinense of the basal eudicot family Trochodendraceae, the flower primordium, together with the much retarded floral subtending bract primordium appear to form a common primordium. The four tepals and the four stamens are initiated in four distinct alternating pairs, the first tepal pair is in transverse position. The four carpels arise in a whorl and alternate with the stamens. This developmental pattern supports the interpretation of the flower as dimerous in the perianth and androecium, but tetramerous in the gynoecium. There is a relatively long temporal gap between the initiation of the stamens and the carpels. The carpel primordia are then squeezed into the narrow gaps between the four stamens. In contrast to Trochodendron, the residual floral apex after carpel formation is inconspicuous. In their distinct developmental dimery including four tepals and four stamens, flowers of Tetracentron are reminiscent of other, related basal eudicots, such as Buxaceae and Proteaceae.     

Flower structure and potential bisexuality in Freycinetia reineckei (Pandanaceae), a species of the Samoa Islands

In Freycinetia reineckei the staminate flower (on the staminate spikes) comprises 3 or 4 (sometimes 2) stamens and a pistillode with 2 (sometimes 4) carpellodes, and the pistillate flower (on the pistillate spikes) is formed of a pistil with 2 (sometimes 4) carpels and of 3 or 4 (sometimes 2) staminodes. This perfect floral homology, also observed in all the other species that were studied with both pistillate and staminate material, strongly suggests that the flower of Freycinetia is basically and potentially bisexual, and may explain the occasional sexual lability and bisexuality of that flower (occurrence of both pistillate and staminate inflorescences, and/or of bisexual inflorescences with bisexual flowers and/or unisexual flowers, on the same individuals) in some species, and also the frequent occurrence of bisexual spikes in this species. These may be partitioned into pistillate, staminate, mixed and sterile zones. In the pistillate zones the flowers have the same aspect and structure as the pistillate flowers. In the staminate zones the flowers generally comprise 3 or 4 (sometimes 2) stamens and a ‘semi-pistil’ some have both stamens and staminodes. The semi-pistils are intermediate between pistils and pistillodes in length, aspect and structure, but always have placentas and ovules. In the mixed zones the flowers are generally formed of a pistil and 3 or 4 (sometimes 2) stamens, and are therefore true hermaphrodite flowers; some have both stamens and staminodes. In the sterile zones the flowers comprise a semi-pistil and 3 or 4 (sometimes 2) staminodes. The staminodes are anatomically very similar to the stamens, especially in the staminate, mixed, and sterile zones, in which they exhibit a wide range of variation in length, aspect and structure. The perfect floral homology as generic character on one hand, and the occasional bisexuality both with and without bisexual flowers and other aspects of sex expression (e.g. occurrence of both pistillate and staminate shoots on the same individuals) in some species on the other hand, seem to indicate that Freycinetia is a basically monoecious, sex changing genus.     

Role of ABA in stamen and pistil development in the normal and solanifolia mutant of tomato (Lycopersicon esculentum)

Summary The role of abscisic acid (ABA) in stamen and pistil development of the normal and solanifolia (sf/sf) mutant of tomato (Lycopersicon esculentum Mill.) was analyzed. The solanifolia mutant produces flowers with separate floral organs, unlike the fused organs of normal flowers, and has greater number of carpels and locules per ovary than the normal. Applications of 10^–5 M ABA to normal floral buds produced flowers with separate stamens, but higher concentrations (10^–4 M ABA) resulted in the complete suppression of stamen growth or stamens that were devoid of anthers. ABA at both 10^–4 and 10^–5 M also induced an increase in the number of carpels and locules in normal flowers, but not in mutant ones. Analysis of endogenous ABA by a radioimmunoassay revealed that the pistils of mutant flowers contained a significantly higher level of ABA than those of normal flowers, but there was no difference in the ABA content of the stamens. The non-fusion of the stamens and the high number of carpels and locules in solanifolia mutant flowers may be explained by the high level of ABA in the floral apex during the initiation and development of carpels.     

Chloranthaceous floral structures from the Late Cretaceous of Sweden

New chloranthaceous floral structures from the Late Cretaceous (Late Santonian/Early Campanian) of Scania, southern Sweden, have provided important new information on theChloranthistemon plants. The material includes well preserved fragments of inflorescence axes with flowers in situ documenting thatChloranthistemon flowers were bisexual and closely resembled those of extantChloranthus (Chloranthaceae). An emended diagnosis is given for the type species of the genus,Chloranthistemon endressii, and a new species,C. alatus, is described. The flowers ofChloranthistemon are small, perianthless and strongly zygomorphic, consisting of a tripartite and broadened androecium borne in an abaxial to lateral position on the monocarpellate ovary, and arranged in the axils of decussate bracts. Stamens are either completely free (C. alatus), or free at the base and coherent at the apex (C. endressii). The apical connective is extensive in both species; elaborated into conspicuous wing-like structures inC. alatus, or into a massive and shield-like structure inC. endressii. Pollen grains ofC. endressii are spheroidal, and reticulate and spiraperturate, while those ofC. alatus are ellipsoidal, tectate and foveolate with a unique combination of a distal colpus and a proximal furrow (colpus?) perpendicular to each other. Ovaries observed in well preserved flowers of both species are small and undifferentiated. Larger, dispersed fruits of chloranthaceous affinity are abundant and distinct, and probably represent at least two or three species, but cannot be linked with certainty to any of theChloranthistemon species described here.     

RFLP mapping of the barley homeotic mutant lax-a

The lax-a homeotic mutant of barley has flowers in which lodicules are replaced by stamens (giving five stamens per flower). RFLP mapping of an F₂ population from a Bonus lax-a ¹ x H. spontaneum cross showed that the mutation was on the short arm of chromosome 7(5H), closely linked to the centromere. An additional F₂ population was used to show that the lax-a mutation gave the five-stamen phenotype in all flowers of 6-rowed spikes and that hoods were elevated and reduced in size in lax-a/Hooded double-mutant plants.