The arrest of development of abortive reproductive organs in the unisexual flower of<Emphasis Type="Italic"> Vitis vinifera</Emphasis> ssp.<Emphasis Type="Italic"> silvestris</Emphasis>

During the first stages of development, flowers of most dioecious species are hermaphroditic, with their transition to unisexual flowers being the result of the developmental arrest of one set of reproductive organs. In this work, we describe the development of male and female flowers of the dioecious wild grape species Vitis vinifera ssp. silvestris through scanning electron microscopy analysis and cytological observations, focusing our attention on the transition from bisexual to unisexual development. We divide floral development of the wild grape into eight stages. Differences between male and female flowers appear first at stage 6, when the style and stigma start to differentiate in female but not in male flowers. Cytological analysis of the slowly growing abortive pistil of male flowers shows that megagametophyte formation is, surprisingly, not inhibited. Instead of pistil abortion in the male flower, sexual determination is accomplished through programmed death of external nucellus cells and some layers of integumentary cells. Sterility of male structures in female flowers follows a different pattern, with microspore abnormalities evident from the time of their release from the tetrad. Sterile microspores and pollen grains in female flowers display an abnormal round shape, lacking colpi and possessing uniformly thickened cell walls that impede germination.     

Anatomy and ontogeny of unisexual flowers in dioecious Woonyoungia septentrionalis （Dandy） Law （Magnoliaceae）

Woonyoungia septentrionalis （Dandy） Law is aceae. The floral morphology and structure of the species a dioecious species with unisexual flowers in Magnoliare conspicuously different from other species and are important to the study of floral phylogeny in this family. The floral anatomy and ontogeny were investigated to evaluate the systematic position of W. septentrionalis, using scanning electron microscopy and light microscopy. All of the floral organs are initiated acropetally and spirally. The carpels are of conduplicated type without the differentiation of stigma and style. The degenerated stamens in the female flowers have the same structures as the normal stamens at the earlier developmental stages, but they do not undergo successive development and eventually degenerate. The male floral apex was observed to have the remnants of carpels in a few investigated samples. As the bisexual flower features could be traced both in the male and female flowers in W. septentrionalis, it suggests that the flower sex in Magnoliaceae tends toward unisexual. As well as the unisexual flowers, the reduced tepals and carpels and concrescence of carpels conform to the specialized tendency in Magnoliaceae, which confirms the derived position of W. septentrionalis in this family. As the initiation pattern of floral parts of W. septentrionalis is very similar to other species in this family, it needs further investigation and especially comparison with species in Kmeria to evaluate the separation of Woonyoungia.     

Development of Sexual Organs in Taihangia rupestris--Different Temperature Requirments for Both Sexual Organ Development in a Bisexual Flower

Optimal environmental temperature for differentiation and development of stamen and pistil have been tested cultured in vitro flowers of Taihangia rupestris. The optimal temperatures for stamen development was quite different from that for pistil development. The most suitable temperature scope for stamen development was on the low verge of 1 ～ 6 'C. Temperature over 18 ℃ led to abortion of all stamens. The temperature scope appropriate for pistil development was near to 6～26 ℃. That below 3 ℃ brought about cessation of most pistil development at early stage. The course of stamen abortion induced by high temperature started from the primordium stage in the stamen development and once this occurred it became irreversible despite of lowering the temperature. It is concluded that the temperature for unisexual male flower, unisexual female flowers and bisexual flowers development should be contralled at 1～3 ℃, 6～26 ℃ and 6～12 ℃ respectively. The results of the present experiment may open up a new train of thoughts for studies on the control of sexual organ development and male sterility.     

Changes in polyamine pattern mediates sex differentiation and unisexual flower development in monoecious cucumber (Cucumis sativus L.)

Changes in the levels of polyamines are associated with fundamental physiological processes such as embryogenesis, induction of flowering, fruit development and ripening, senescence, and responses to environmental stresses, but the role of polyamines in sex differentiation and unisexual flower development has not been deeply studied. To extend the knowledge on the regulatory mechanisms of flowering in monoecious plant (producing unisexual flowers), we investigated the morphogenesis and free polyamine levels in Cucumis sativus during sex differentiation and unisexual flower development in vitro using histocytological and biochemical methods. As shown in our study, floral development in vitro was undisturbed and flowers of both sexes were produced. Sex differentiation relied on preventing the development of generative organs of the opposite sex, as we observed carpel repression in male flowers and stamen repression in female flowers. Pollen viability was negatively correlated with female flower development on the same node. Biochemical analysis revealed increased accumulation of aliphatic amines (tri, tetra‐amines) in generative (flower buds and flowers) compare to vegetative (axillary buds and leaves) organs. Undifferentiated floral buds contained elevated levels of agmatine, cadaverine, spermidine and spermine. Sex differentiation was associated with significantly decreased levels of agmatine and cadaverine. Our results showed that female flowers contained higher levels of total polyamine than male flowers. The increased level of cadaverine was associated with macrogametogenesis and female flower maturation. Putrescine was important for male flower development. Such results support the hypothesis that aliphatic amines are involved in unisexual flower development.     

Characterization of unisexual flower development in the endangered mahogany tree Swietenia macrophylla King. (Meliaceae)

The selection of candidate plus trees of desirable phenotypes from tropical forest trees and the rapid devastation of the natural environments in which these trees are found have created the need for a more detailed knowledge of the floral and reproductive biology of tropical tree species. In this article, the organogenic processes related to unisexual flower development in tropical mahogany, Swietenia macrophylla , are described. Mahogany inflorescences at different developmental stages were evaluated using scanning electron microscopy or optical microscopy of histological sections. The unisexual flowers of S. macrophylla are usually formed in a thyrse, in which the positions of the female and male flowers are not random. Differences between male and female flowers arise late during development. Both female and male flowers can only be structurally distinguished after stage 9, where ovule primordia development is arrested in male flowers and microspore development is aborted in female flower anthers. After this stage, male and female flowers can be distinguished by the naked eye as a result of differences in the dimensions of the gynoecium. The floral characteristics of S. macrophylla (distribution of male and female flowers within the inflorescence, and the relative number of male to female flowers) have practical implications for conservation strategies of this endangered species.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 529–535.     

Pollination adaptations of group-by-group stamen movement in a meadow plant with temporal floral closure

Floral sexual organ (stamen and pistil) movements are selective adaptations that have different functions in male-female reproduction and the evolution of flowering plants. However, the significance of stamen movements in the spatial–temporal function and separation of male and female organs has not been experimentally determined in species exhibiting floral temporal closure. The current study investigated the role of slow stamen (group-by-group) movement in male-female sexual function, and the effect of stamen movement on pollen removal, male-male and male-female interference, and mating patterns of Geranium pratense, a plant with temporal floral closure. This species uses stamen group-by-group movement and therefore anther-stigma spatial–temporal separation. Spatial separation (two whorls of stamen and pistil length) was shown to be stronger than temporal separation. We found that stamen movements to the center of the flower increase pollen removal, and the most common pollinators visited more frequently and for longer durations during the male floral stage than during the female floral stage. Petal movements increased both self-pollen deposition rate and sexual interference in G. pratense. The fruit and seed set of naturally and outcrossed pollinated flowers were more prolific than those of self-pollinated flowers. Group-by-group stamen movement, dehiscence of stamens, pistil movement, and male-female spatial–temporal functional separation of G. pratense before floral temporal closure may prevent male-female and stamen-stamen interference and pollen discounting, and may increase pollen removal and cross-pollination.     

Multiple developmental processes underlie sex differentiation in angiosperms

The production of unisexual flowers has evolved numerous times in dioecious and monoecious plant taxa. Based on repeated evolutionary origins, a great variety of developmental and genetic mechanisms underlying unisexual flower development is predicted. Here, we comprehensively review the modes of development of unisexual flowers, test potential correlations with sexual system, and end with a synthesis of the genetics and hormonal regulation of plant sex determination. We find that the stage of organ abortion in male and female flowers is temporally correlated within species and also confirm that the arrest of development does not tend to occur preferentially at a particular stage, or via a common process.     

栝楼不同性别花芽分化形态解剖特征观察

采用体视显微镜、石蜡切片和树脂切片技术对栝楼(Trichosanthes kirilowii Maxim.)不同性别花芽分化发育时期的外部形态和内部解剖结构进行了观察。结果显示,栝楼花为雌雄异株,仅有雌花、雄花两种性别分化,且雄花的发育速度明显快于雌花的发育速度。栝楼雌雄花芽长0.2 mm左右已完成性别分化;栝楼雄花为单性花,分化过程可分为6个时期,整个发育过程仅见雄蕊原基的分化及生长。栝楼雌花为"两性花",分化过程可分为7个时期,存在雌蕊和雄蕊共同发育阶段,后期雄蕊发育败退。本研究明确了不同性别栝楼花芽发育发生的各个阶段、形态变化特点、外部形态变化特征以及雌雄花芽的分化差异,建立了雌雄花芽内部结构分化与外部形态之间相关性,为栝楼早期幼苗鉴定及性别分化研究提供了一定的参考。     

Isolation of a partial sequence of a putative nucleotide sugar epimerase, which may involve in stamen development in cucumber (Cucumis sativus L.)

Sex determination is the most widely studied subject in cucumber. The sex of cucumber plants can be monoecious, hermaphrodite, gynoecious, androecious, or andromonoecious. Besides environmental factors, three major genes, F/f, M/m, and A/a mainly govern the sex types in cucumber. Regardless of their sex all floral buds are bisexual at the early bud stage. A stage specific arrest of either stamen or carpel leads to unisexual flower development. The possible downstream product of the interaction of the sex determining genes that may directly allow the growth or selectively arrest stamen or pistil is not yet identified. Therefore, in the current study, we performed suppression subtractive hybridization using floral buds from nearly isogenic gynoecious and hermaphrodite cucumber plants and identified for the first time a cDNA homologous to nucleotide sugar epimerase. The expression level of the isolated putative nucleotide sugar epimerase is weak in female floral buds but strong in bisexual and male flowers. The weak level of the putative nucleotide sugar epimerase may be an indication for its improper functioning, which may influence stamen development in cucumber plants.     

Unisexual flower, spikelet, and inflorescence development in monoecious/dioecious Bouteloua dimorpha (Poaceae, Chloridoideae)

Unisexual flowers have evolved repeatedly in the angiosperms. In Poaceae, multiple transitions from bisexual to unisexual flowers are hypothesized. There appear to be at least three distinct developmental mechanisms for unisexual flower formation as found in members of three subfamilies (Ehrhartoideae, Panicoideae, Pharoideae). In this study, unisexual flower development is described for the first time in subfamily Chloridoideae, as exemplified by Bouteloua dimorpha. Scanning electron microscopy (SEM) and anatomy were used to characterize the development of male (staminate) and female (pistillate) flowers, spikelets, and inflorescences. We found the developmental pathway for staminate flowers in B. dimorpha to be distinct from that described in the other three subfamilies, showing gynoecial arrest occurs at a different stage with possible loss of some cellular contents. However, pistillate flowers of B. dimorpha had some similarity to those described in other unisexual-flowered grasses, with filament and anther differentiation in abortive stamens. Comparing our findings with previous reports, unisexual flowers seem to have evolved independently in the four examined grass subfamilies. This analysis suggests the action of different genetic mechanisms, which are consistent with previous observations that floral unisexuality is a homoplasious condition in angiosperms.     

Architectural effects mimic floral sexual dimorphism in Solanum (Solanaceae)

Factors underlying apparent floral sexual dimorphism were examined in six species of andromonoecious Solanum section Lasiocarpa (Solanaceae). Both multivariate and univariate analyses show that hermaphroditic flowers are significantly larger than staminate flowers for all features measured. Thus, flowers could be characterized as sexually size dimorphic. However, when size variation due to flower position (architecture) is controlled experimentally, differences between the floral genders for the nongynoecial characters disappear; there is no difference in corolla or androecium size. Staminate flowers appear to be generally smaller than hermaphroditic flowers, not because of any difference related to primary sexual function, but because they tend to occur in the distal regions of each inflorescence. In contrast, significant differences between hermaphroditic and staminate flowers for primary female traits (ovary, style, and stigma) remain after controlling for position: the two floral types are truly dimorphic for these characters. We show that consideration of architectural effects can direct and refine hypotheses concerning the evolution of andromonoecy. More generally, if architectural effects on flower size are common among taxa with unisexual flowers, then these effects may contribute to the common perception of size dimorphism in taxa with unisexual flowers.     

Pollinator responses to variation in floral display and flower size in dioecious Sagittaria latifolia (Alismataceae)

In animal-pollinated plants with unisexual flowers, sexual dimorphism in floral traits may be the consequence of pollinator-mediated selection. Experimental investigations of the effects of variation in flower size and floral display on pollinator visitation can provide insights into the evolution of floral dimorphism in dioecious plants. Here, we investigated pollinator responses to experimental arrays of dioecious Sagittaria latifolia in which we manipulated floral display and flower size. We also examined whether there were changes in pollinator visitation with increasing dimorphism in flower size. In S. latifolia, males have larger flowers and smaller floral displays than females. Visitation by pollinators, mainly flies and bees, was more frequent for male than for female inflorescences and increased with increasing flower size, regardless of sex. The number of insect visits per flower decreased with increasing floral display in males but remained constant in females. Greater sexual dimorphism in flower size increased visits to male inflorescences but had no influence on the number of visits to female inflorescences. These results suggest that larger flower sizes would be advantageous to both females and males, and no evidence was found that females suffer from increased flower-size dimorphism. Small daily floral displays may benefit males by allowing extended flowering periods and greater opportunities for effective pollen dispersal.     

Functional analysis of synchronous dichogamy in flowering rush, Butomus umbellatus (Butomaceae)

Dichogamy is one of the most widespread floral mechanisms in flowering plants and is thought to have evolved to reduce interference between pollen import and export within flowers, especially self-pollination. Self-pollination between flowers may also be reduced if dichogamy is synchronous among flowers on an inflorescence. The analysis of dichogamy at both levels requires that the sexual phases of individual flowers be defined functionally in terms of pollen deposition and removal. We conducted morphological and functional analyses to investigate the degree of dichogamy within flowers and the synchronicity of dichogamy between flowers within inflorescences in an emergent, aquatic monocot, flowering rush (Butomus umbellatus). Based on daily observations of the development of marked flowers, data on the schedule of anther dehiscence within flowers, and repeat surveys of floral sex ratios in three populations, individual flowers appear to be strictly protandrous. On average, each flower spends ～1 d in each of male and female phases with an intervening 1-d neuter phase during which there is no available pollen in anthers and stigmas are not yet exposed to receive pollen. Morphological criteria used to delimit the beginning and end of each of these three sex phases were validated by quantifying the temporal schedule of pollen removal from anthers and pollen deposition on stigmas. Experimental pollinations showed that the morphological changes marking the end of female phase are hastened by pollen deposition. At the umbel level, synchronous development within sequential cohorts of flowers reduced overlap of male and female sexual phases between flowers. On average (±1 SE), 72 ± 3% of flowers completed their female phase while no other flowers on the same umbel were in male phase. Computer simulations of umbel development showed that this value is significantly higher than expected if the timing of flower development within umbels was random (30 ± 1%). Surveys of floral sex ratios in three populations revealed that 87% of umbels were either unisexual male or female at any given time. Pollinators usually visited more than one flower in sequence when foraging on umbels, suggesting that synchronous dichogamy may be an adaptation to avoid geitonogamy. The adaptiveness of both flower- and umbel-level dichogamy is also suggested because both traits are expressed to a lesser extent in obligately clonal, triploid populations, where flowers do not make seeds and hence floral adaptations are not maintained by natural selection.     

Homeosis, morphogenetic gradient and the determination of floral identity in the inflorescences ofPhilodendron solimoesense (Araceae)

A comparative developmental study of the inflorescence ofPhilodendron solimoesense was conducted using scanning electron microscopy. The spadix ofP. solimoesense is characterized by unisexual flowers. Staminate flowers are initiated on the upper portion of the spadix while pistillate flowers develop on the lower portion of the spadix. An intermediate zone located between the upper male and lower female portion of the inflorescence consists of sterile male flowers. Within this intermediate zone a row of flowers exhibit polarity with respect to the identity of sexual organs. Stamens are initiated on the flank of the floral meristem facing the upper male zone and carpels are initiated on the portion of the floral meristem facing the lower female zone. The resulting flowers therefore assume a bisexual identity. At the level of the inflorescence, all floral buds are initiated along a series of contact parastichies and the continuity of these parastichies is not disrupted at any level in the male, intermediate, and female zones on the spadix. Results from this study support the presence of a morphogenetic gradient acting at the level of the inflorescence and appears to be independent of the boundaries of floral primordia.     

Scanning Electron Microscopic Studies on the Development of the Organs of Litchi Flower

The pattern of development of the floral parts of litchi (Litchi chinensis Sonn.) flower was followed using scanning electron microscopy. Before making scanning electron microscopic observations, specimens were tannin-osmium impregnated and critical point dried. In the bisexual flower, floral organogenesis starts with the formation of protrusions near the floral apex. The two to three protrusions present at the apical region of the floral apex later expand and fuse to form the ovary. At the upper middle region of the ovary another protrusion develops which later becomes the style and the stigma. When the flower matures the tip of the style not only splits but also becomes twisted. On the upper side of the stigma there are numerous papillate cells. These cells are covered with mucilage when fully mature. The development of the filament and anther begins a little bit earlier than the gynoecium. The first sign of androecium development begins when protrusions start to develop around the floral apex. Each litchi flower possesses 6 to 10 anthers. In addition to forming bisexual flowers, litchi also produces a large number of male and female unisexal flowers. But under the scanning electron microscope it is very difficult to distinguish accurately between male and female flowers, because both flowers invariably give rise to some poorly developed organs of the opposite sex. Thus it seems that all flowers in litchi are potentially bisexual and only at the final stage of development (i.e. about 50 days after floral initiation) sex organs fail to develop properly in some flowers.     

Developmental analyses reveal early arrests of the spore-bearing parts of reproductive organs in unisexual flowers of cucumber (<Emphasis Type="Italic">Cucumis sativus</Emphasis> L.)

To understand the regulatory mechanisms governing unisexual flower development in cucumber, we conducted a systematic morphogenetic analysis of male and female flower development, examined the dynamic changes in expression of the C-class floral organ identity gene CUM1, and assessed the extent of DNA damage in inappropriate carpels of male flowers. Accordingly, based on the occurrence of distinct morphological events, we divided the floral development into 12 stages ranging from floral meristem initiation to anthesis. As a result of our investigation we found that the arrest of stamen development in female flowers, which occurs just after the differentiation between the anther and filament, is mainly restricted to the primordial anther, and that it is coincident with down-regulation of CUM1 gene expression. In contrast, the arrest of carpel development in the male flowers occurs prior to the differentiation between the stigma and ovary, given that no indication of ovary differentiation was observed even though CUM1 gene expression remained detectable throughout the development of the stigma-like structures. Although the male and female reproductive organs have distinctive characteristics in terms of organ differentiation, there are two common features regarding organ arrest. The first is that the arrest of the inappropriate organ does not affect the entirety of the organ uniformly but occurs only in portions of the organs. The second feature is that all the arrested portions in both reproductive organs are spore-bearing parts.Abbreviations SEM Scanning electron microscopy - TEM Transmission electron microscopy - TUNEL TdT-mediated dUTP nick-end labeling     

BREEDING SYSTEM IN FICUS CARICA,THE COMMON FIG. I. FLORAL DIVERSITY

Coevolution in Ficus carica (Moraceae) and the fig wasp (Blastophaga psenes, Agaonidae, Chalcidoidea) has resulted in a complex breeding system involving two tree morphs (Caprifig and Edible fig), three floral forms (long-styled female, short-styled female, and male flowers) and the insect pollinator. The two female floral forms have been reported to differ only in style length and stigma shape. In the present study, we demonstrate that the two female flowers differ from inception—short-styled flower primordia are smaller and exhibit significantly greater individual variation than do those of the long-styled flower, and the relative growth rate of each flower type differs. Mature forms exhibit disparity in style length, in stigma characteristics, and in degree of fusion of stylar lobes. Female flowers of both tree morphs are unisexual from inception. Male flowers of the Caprifig tree morph are initiated as hermaphrodites and gynoecium abortion occurs before megaspore mother cell stage. A single inflorescence therefore expresses two pathways to unisexuality. Hermaphrodite flower primordia were repeatedly found in the supposedly unisexual female syconium of the Edible fig tree morph. Based on its developmental morphology, Ficus carica appears to be of gynomonoecious ancestry.     

DEVELOPMENT AND VASCULATURE OF THE FLOWERS OF LOPHOTOCARPUS CALYCINUS AND SAGITTARIA LATIFOLIA (ALISMACEAE)

The development of the bisexual flower of Lophotocarpus calycinus and of the unisexual flowers of Sagittaria latifolia has been observed. In all eases floral organs arise in acropetal succession. In L. calycinus, after initiation of the perianth, the first whorl of stamens to form consists of six stamens and is ordinarily followed by two alternating whorls of six stamens each. The very numerous carpels arc initiated spirally. In the male flower of S. latifolia the androecium develops in spiral order. A few rudimentary carpels appear near the floral apex after initiation of the stamens. There are no staminodia. The female flower has a similar developmental pattern to that of Lophotocarpus except that a prominent residual floral apex is left bare of carpels. The vascular system in all flowers is semiopen, with vascular bundles passing to the floral organs in a pattern unrelated to the relative positions of those organs. The androecia of these two taxa are similar to those of some Butomaceae and relationships based on ontogeny and morphology are suggested. The gynoecia are meristically less specialized but morphologically more specialized than the gynoecia of Butomaceae.     

The evolution of unisexual flowers: morphological and functional convergence results from diverse developmental transitions

Unisexual flower morphology was examined within a phylogenetic context in order to identify developmental transitions associated with the multiple origins of dioecy in flowering plants. Historically, two categories of unisexual flowers have been recognized: type I flowers exhibit rudiments of the nonfunctional organ type, while type II flowers bear no vestigial sexual organs. Mapping of these flower types onto a composite phylogeny shows that type II morphology is homoplasious and has resulted from at least four distinct evolutionary developmental pathways. The historical assignment of unisexual flowers into only two morphological types has masked important developmental and evolutionary dynamics.