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Chrysanthemum morifolium is a gynomonoecious plant that bears both female zygomorphic ray florets and bisexual actinomorphic disc florets in the inflorescence. This sexual system is quite prevalent in Asteraceae, but poorly understood. CYCLOIDEA (CYC) 2 subclade transcription factors, key regulators of flower symmetry and floret identity in Asteraceae, have also been speculated to function in reproductive organs and could be an entry point for studying gynomonoecy. However, the molecular mechanism is still unclear. On the other hand, the Arabidopsis WUSCHEL (WUS) transcription factor has been proven to play a vital role in the development of reproductive organs. Here, a WUS homologue (CmWUS) in C. morifolium was isolated and characterized. Overexpression of CmWUS in A. thaliana led to shorter siliques and fewer stamens, which was similar to CYC2-like genes reported before. In addition, both CmWUS and CmCYC2 were highly expressed in flower buds during floral organ differentiation and in the reproductive organs at later development stages, indicating their involvement in the development of reproductive organs. Moreover, CmWUS could directly interact with CmCYC2d. Thus, our data suggest a collaboration between CmWUS and CmCYC2 in the regulation of reproductive organ development in chrysanthemum and will contribute to a further understanding of the gynomonoecious sexual system in Asteraceae.

PREMISE OF THE STUDY: Abiotic environmental factors are often considered to be important in the distribution and maintenance of variation in sexual systems in flowering plants. Associations between sexes and abiotic factors are well documented in dioecious systems, but much less is known about this relationship in other sexually polymorphic systems. Species that are highly variable in sexual expression and habitat distribution can provide insights into the role of abiotic factors in maintaining variation in sexual expression. METHODS: Focusing on a sexually polymorphic species, Geranium transversale, we measured sexual expression at both the flower and the plant level and examined vegetative and floral traits, pollen deposition, and reproductive success. We also tested for correlations between sexual expression and other traits and examined whether and how these traits covaried with abiotic environmental conditions. KEY RESULTS: We identified unique variation of sexual expression in G. transversale. There are four sexual morphs that display different combinations of the three flower types (pistillate, staminate, and perfect). Sexual morphs that are phenotypically more female (i.e., female and gynomonoecious morphs) are found in wetter and milder environments, and flower earlier than morphs that are more male (i.e., hermaphroditic and andromonoecious morphs). Additionally, floral organ size and reproductive success are influenced not only by the flower type but also by the sexual morph of the plant. CONCLUSIONS: Environmental conditions are likely to cause some of the variation in sexual expression found in G. transversale. Both genetic and ecological factors likely contribute to the maintenance of sexual variation in this species.

• A range of hypothesized evolutionary pathways has been proposed for describing the evolution of dioecy. However, the evolutionary links between other sexual systems not directly involved in dioecy evolution remain largely unexplored, and hence, a comprehensive picture of evolutionary transitions between sexual systems is still lacking. • Here, we explored the diversity and evolution of sexual systems in Asteraceae, the largest family of flowering plants, where almost all sexual systems are present. We used a phylogenetic approach to build a model of evolutionary transitions between sexual systems. • The best model involved nine transitions, including those from hermaphroditism to andromonoecy, gynomonoecy and gynodioecy, those from gynomonoecy to monoecy and trimonoecy, two transitions to dioecy -one through gynodioecy and the other through monoecy - and reversals from monoecy to gynomonoecy and from gynomonoecy to hermaphroditism. • Our reconstruction of the evolution of sexual systems in Asteraceae provided, for the first time, a joint view of the evolutionary transitions between seven sexual systems, unveiling the evolutionary links between monomorphic sexual systems. A pathway from hermaphroditism to monoecy through gynomonoecy, instead of from andromonoecy, was highly supported, which was consistent with a gradient of floral gender specialization.

Most orchid species exhibit an extreme case of hermaphroditism, owing to the fusion of male and female organs into a gynostemium. Exceptions to this rule have only been reported from the subtribes Catasetinae and Satyriinae. Here, I report an additional orchidaceous example whose flowers are not always hermaphroditic. In several Japanese populations of Eulophia zollingeri (Rchb.f.) J.J.Sm, a widespread Asian and Oceanian orchid, some flowers possess both the anther (i.e., anther cap and pollinaria) and stigma, whereas others possess only the stigma. Therefore, pollination experiments, an investigation of floral morphology and observations of floral visitors were conducted to understand the reproductive biology of E. zollingeri in Miyazaki Prefecture, Japan. It was confirmed that E. zollingeri studied here possesses a gynomonoecious reproductive system, a sexual system in which a single plant has both female flowers and hermaphroditic flowers. In addition, hermaphroditic flowers often possess an effective self-pollination system while female flowers could avoid autogamy but suffered from severe pollinator limitation, due to a lack of agamospermy and low insect-mediated pollination. The present study represents the first documented example of gynomonoecy within Orchidaceae. Gynomonoecy in E. zollingeri may be maintained by the tradeoff in reproductive traits between female flowers (with low fruit set but potential outcrossing benefits) and hermaphroditic flowers (with high fruit set but inbreeding depression in selfed offspring). This mixed mating is probably important in mycoheterotrophic E. zollingeri because it occurs in shaded forest understorey with a paucity of pollinators.

PREMISE OF THE STUDY: It has been hypothesized that two flower types permit flexible allocation of resources to female and male functions, yet empirical evidence for the sex‐allocation hypothesis remains scarce in gynomonoecious species. To characterize resource allocation to pistillate and perfect flowers and allocation of perfect flowers between gynomonoecious and hermaphroditic individuals, we examined the flexibility and whether female‐biased allocation increases with plant size in the hermaphroditic‐gynomonoecious herb Eremurus anisopterus. METHODS: Frequency of gynomonoecious individuals, flower production, and plant size were investigated in different populations. Floral allocation was compared among the three flower types of E. anisopterus. KEY RESULTS: Frequency of gynomonoecious plants varied from 2–17% in nine populations. Only larger plants produced female flowers at the bottom of racemes. Both female and perfect flower production tended to increase proportionately with plant size in gynomonoecious individuals. Female flowers did not produce less biomass than perfect flowers from hermaphroditic or gynomonoecious plants. However, both female and perfect flowers from gynomonoecious individuals had lighter stamen mass, but larger pistil mass, than perfect flowers from hermaphrodites. CONCLUSIONS: Although the prediction of an increase in female flower number with plant size was not observed in E. anisopterus, the flexibility of sex allocation in gynomonoecious species was confirmed in that gynomonoecious individuals had a female‐biased floral allocation compared to hermaphroditic individuals. Such comparisons of gynomonoecious to hermaphroditic individuals permit us to unveil a sexual adjustment strategy: flexibility of sexual investments within plants.

Predominantly outcrossing plant species are expected to accumulate recessive deleterious mutations, which can be purged when in a homozygous state following selfing. Individuals may vary in their genetic load because of different selfing histories, which could lead to differences in inbreeding depression among families. Lineage-dependent inbreeding depression can appear in gynodioecious species if obligatory outcrossed females are more likely to produce female offspring and if partially selfing hermaphrodites are more likely to produce hermaphrodites. We investigated inbreeding depression at the zygote, seed, and germination stages in the gynomonoecious-gynodioecious Dianthus sylvestris, including pure-sexed plants and a mixed morph. We performed hand-pollinations on 56 plants, belonging to the three morphs, each receiving 2-3 cross treatments (out-, sib- and self-pollination) on multiple flowers. Effects of cross treatments varied among stages and influenced seed provisioning, with sibling competition mainly occurring within outcrossed fruits. We found significant inbreeding depression for seed mass and germination and cumulative early inbreeding depression varied greatly among families. Among sex morphs, we found that females and hermaphrodites differed in biparental inbreeding depression, whereas uniparental was similar for all. Significant inbreeding depression levels may play a role in female maintenance in this species, and individual variation in association with sex-lineages proclivity is discussed.

• Premise of the study: In heterocarpy, fruits with different morphologies have been associated with alternative strategies of dispersal, germination, dormancy, and seedling competitive ability. In heterocarpic species, it is common to find fruits with competitive or dispersal syndromes. The competitive advantage of nondispersing fruits has been frequently attributed to their larger size, but recent studies have suggested that this could also be mediated by germination time. The main objective of our study was to investigate which factor, fruit type or germination time, most affects plant performance and, consequently, competitive ability, using the heterocarpic species Anacyclus clavatus.• Methods: To explore the effects of achene type and germination time on plant performance, we followed an innovative experimental approach including two experiments: one allowing for differences in germination time, and the other evaluating the effect of achene type alone by synchronizing germination time.• Key results: A significant effect of germination time on several postdispersal life-history traits was observed: Achenes that germinated earlier produced plants with higher biomass and reproductive effort. When germination time was controlled, no significant differences were observed in any of the traits.• Conclusions: The competitive advantage of achenes with different morphologies was mainly mediated by germination time and not by differences in size or other intrinsic traits. The consequences of these results are discussed in light of the dispersal–competition trade-off. Our experimental approach (i.e., the synchronization of germination time) revealed the importance of manipulative experiments for testing the effects of germination time on plant survival and performance.

Background Coccinia grandis is a dioecious species of Cucurbitaceae having heteromorphic sex chromosomes. The chromosome constitution of male and female plants is 22 + XY and 22 + XX respectively. Y chromosome of male sex is conspicuously large and plays a decisive role in determining maleness. Sex modification has been studied in hypogynous Silene latifolia (Caryophyllaceae) but there is no such report in epigynous Coccinia grandis . Moreover, the role of organ identity genes during sex expression in Coccinia has not been evaluated earlier. Investigations on sexual phenotypes of C. grandis including a rare gynomonoecious (GyM) form and AgNO 3 mediated sex modification have added a new dimension to the understanding of sex expression in dioecious flowering plants. Results Morphometric analysis showed the presence of staminodes in pistillate flowers and histological study revealed the absence of carpel initials in male flowers. Though GyM plant had XX sex chromosomes, the development of stamens occurred in hermaphrodite flowers but the pollens were not fertile. Silver nitrate (AgNO 3 ) application enhanced stamen growth in wild type female flowers like that of GyM plant but here also the pollens were sterile. Differential expression of CgPI could be involved in the development of different floral phenotypes. Conclusions The three principle factors, Gynoecium Suppression (Su F ), Stamen Promoting Factor (SPF) and Male Fertility (m F ) that control sex expression in dioecious C. grandis assumed to be located on Y chromosome, play a decisive role in determining maleness. However, the characteristic development of stamens in hermaphrodite flowers of GyM plant having XX sex chromosomes indicates that Y-linked SPF regulatory pathway is somehow bypassed. Our experimental findings together with all other previous chromosomal and molecular cytogenetical data strongly support the view that C. grandis could be used as a potential model system to study sex expression in dioecious flowering plant.

Species and populations can be categorized by their sexual systems, depending on the spatial distribution of female and male reproductive structures within and among plants. Although a high diversity of sexual systems exists in Silene, their relative frequency at the genus and infrageneric level is unknown. Here, we carried out an extensive literature search for direct or indirect descriptions of sexual systems in Silene species. We found descriptions of sexual systems for 98 Silene species, where 63 and 35 correspond to the phylogenetically supported subgenera Silene and Behenantha, respectively. Hermaphroditism was the commonest sexual system (58.2 %), followed by dioecy (14.3 %), gynodioecy (13.3 %) and gynodioecy–gynomonoecy (i.e. hermaphroditic, female and gynomonoecious plants coexisting in the same population; 12.2 %). The presence of these sexual systems in both subgenera suggests their multiple origins. In 17 species, the description of sexual systems varied, and in most cases these differences corresponded to variations within or among populations. Interestingly, the poorly studied gynodioecy–gynomonoecy sexual system showed similar frequency to dioecy and gynodioecy in both subgenera. In addition, the incidence of gynodioecy–gynomonoecy was analysed in the species of section Psammophilae (Silene littorea, S. psammitis, S. adscendens and S. cambessedesii), in a survey of 26 populations across the distribution area of the species. The four species showed gynomonoecy–gynodioecy in most populations. Hermaphrodites were the most frequent morph, with a low number of females and gynomonoecious plants in all populations. The frequency of sexual morphs varied significantly among the studied populations but not among species. Female plants generally produced smaller numbers of flowers than hermaphroditic or gynomonoecious plants, and the percentages of female flowers per population were low. All these findings suggest that the gynodioecious–gynomonoecious sexual system in section Psammophilae is closer to hermaphroditism or gynomonoecy than gynodioecy.

Some species described as gynodioecious are truly gynomonoecious-gynodioecious. Three distinct phenotypes may be found in their natural populations-female and hermaphrodite pure-sexed plants bearing either only pistillate or perfect flowers, respectively, and mixed plants bearing both types of flowers. In one such species, Dianthus sylvestris, we investigated mating system parameters using allozyme data. Outcrossing rates and correlations of outcrossed paternity were calculated for the three types of plants and separately for pistillate and perfect flowers on mixed plants. The mean outcrossing rate for the population was$t_m \\pm SD = 0.885 \\pm 0.032$. Females were more outcrossed than hermaphrodites ($0.987 \\pm 0.112$and$0.790 \\pm 0.076$, respectively), whereas mixed plants were not significantly more or less outcrossed than hermaphrodites ($0.840 \\pm 0.060$). Within mixed plants, perfect flowers showed an intermediate outcrossing rate ($0.898 \\pm 0.057$), whereas pistillate flowers were as selfed as perfect flowers on hermaphrodite plants ($0.782 \\pm 0.111$). Family estimates of outcrossing rates were highly variable. Globally, no biparental inbreeding was detected in this species, and there was a mean of$61.5 \\pm 19.9\\%$of full-sibs within families. Floral dimorphism between small pistillate and large perfect flowers together with pollinator preference for larger flowers could explain the observed patterns for both mating parameters. The advantages of gynomonoecy-gynodioecy are discussed. We conclude that mixed plants do not reduce selfing for all flowers on a plant, but perfect flowers on these plants seem to have an outcrossing advantage.