Floral Symmetry Research Articles

Selection for floral integration and trait variation in zygomorphic flowers of Aconitum japonicum ssp. subcuneatum (Ranunculaceae)

Pollinator-mediated selection might lead to among-trait differences in the degree and pattern of floral integration and intra-flower variation. To examine the patterns of intra-flower variation in floral traits, including nectar volume, we performed a field study using the zygomorphic flowers of Aconitum japonicum ssp. subcuneatum. We investigated (1) correlations between the sizes of the left and right sepals and petals, (2) variation in floral traits among plants, within plants and within flowers, (3) effects of sexual phases on floral integration variation in floral and nectar traits, and (4) the effect of size and intra-flower variation in traits of the left and right sepals and petals on pollen removal by pollinators. Lateral sepal area, but not lower sepal area, was highly correlated between the left and right sepals. Floral traits were more integrated during the male phase than during the female phase. Nectar standing crop in male-phase flowers correlated with helmet height and lateral and lower sepal area, but in female-phase flowers it only correlated with spur length. While intra-flower variance in lateral sepal area accounted for approximately 10% of the overall variance in these traits, the variance in lower sepal area accounted for 70% of the overall variance. Lateral sepal area had a negative effect on the number of pollen grains remaining after pollinator visits. Low variance in lateral sepals within flowers and measurements of pollen removal suggest that lateral sepals play a more important role in pollen export than the other traits. Left and right sepals may be the targets of selection for symmetry in zygomorphic flowers.

Chamaelirium shimentaiense (Melanthiaceae), a new species from Guangdong, China

Chamaelirium shimentaiense (Melanthiaceae), a new species from Shimentai Nature Reserve, Guangdong Province, China is described. This species is similar to C. shiwandashanense and C. viridiflorum in having actinomorphic flowers with six tepals and unilocular anthers, but is easily distinguished from them by its strongly undulate leaf blades with crispulate margins and inconspicuous venation, purplish tepals with a slightly broadened and obtuse apex, and purple‐tinged filaments. The infrageneric assignment of this species is discussed and a color plate is provided.

Nectar robbers deter legitimate pollinators by mutilating flowers

Nectar robbing – harvesting nectar illegitimately – can have a variety of outcomes for plant sexual reproduction and for the pollinator community. Nectar robbers can damage flowers while robbing nectar, which could affect the behavior of subsequent flower visitors and, consequently, plant reproduction. However, only nectar manipulation by nectar robbers has so far received attention. We found a short‐tongued bee, Hoplonomia sp. (Halictidae), mutilating the conspicuous lower petal of the zygomorphic flowers of Leucas aspera (Lamiaceae) while robbing nectar. We hypothesized that the mutilation of the conspicuous lower petal deters legitimate pollinators on L. aspera flowers, which, in turn, might affect plant reproduction. We first assessed the proportion of naturally‐robbed flowers in plant populations for three years to confirm that it was not a purely local phenomenon due to a few individual bees. We then studied diversity, community and visitation characteristics of pollinators, nectar dynamics and fruit set in unrobbed and robbed open flowers in naturally‐robbed populations. The proportion of robbed flowers varied significantly across sites and years. Robbing did not affect nectar dynamics in flowers, but it did alter flower morphology, so much so that it reduced pollinator visitation and altered the pollinator community on robbed flowers. However, the maternal function of plant reproduction was not affected by nectar robbing. This study for the first time shows that a nectar robber can have an ecologically significant impact on floral morphology.

CmCYC2-like transcription factors may interact with each other or bind to the promoter to regulate floral symmetry development in Chrysanthemum morifolium.

The complex capitulum of Chrysanthemum morifolium is often comprised of bilaterally symmetrical ray florets and radially symmetrical disc florets. The TCP transcription factor clade CYCLOIDEA2 (CYC2) appears to play a vital role in determining floral symmetry and in regulating floral organ development in Asteraceae. Our previous study identified six CmCYC2 genes from chrysanthemum and showed that CmCYC2c participated in the regulation of ray floret identity. However, the functions of other CmCYC2 genes and the underlying molecular mechanism of CmCYC2-mediated floral development regulation in chrysanthemums have not been elucidated. In this study, we analysed the function of CmCYC2 genes by ectopic expression of CmCYC2 in Arabidopsis. Then, we examined the protein-protein interaction using yeast two-hybrid (Y2H) and bimolecular fluorescence complementation (BiFC) assays. Finally, we analysed the protein-DNA interaction using yeast one-hybrid (Y1H) and dual-luciferase reporter assays. We found that ectopic expression of CmCYC2 genes in the Arabidopsis tcp1 mutant changed its floral symmetry and flowering time. Y2H and BiFC assays confirmed three pairs of interactions between CmCYC2 proteins, that is, CmCYC2b-CmCYC2d, CmCYC2b-CmCYC2e and CmCYC2c-CmCYC2d, suggesting that heterodimeric complexes may form between CmCYC2 proteins to increase their functional specificity. The results of Y1H and dual-luciferase reporter assays indicate that CmCYC2c can bind to the promoter of ClCYC2f. Our findings provided clues that CmCYC2-like transcription factors may interact with each other or bind to the promoter to regulate floral symmetry development in C. morifolium. KEY MESSAGE: CmCYC2-like transcription factors may interact with each other or bind to the promoter to regulate floral symmetry development in Chrysanthemum morifolium.

A design principle for floral organ number and arrangement in flowers with bilateral symmetry.

The bilateral symmetry of flowers is a striking morphological achievement during floral evolution, providing high adaptation potential for pollinators. The symmetry can appear when floral organ primordia developmentally initiate. Primordia initiation at the ventral and dorsal sides of the floral bud is differentially regulated by several factors, including external organs of the flower and CYCLOIDEA (CYC) gene homologues, which are expressed asymmetrically on the dorso-ventral axis. It remains unclear how these factors control the diversity in the number and bilateral arrangement of floral organs. Here, we propose a mathematical model demonstrating that the relative strength of the dorsal-to-ventral inhibitions and the size of the floral stem cell region (meristem) determines the number and positions of the sepal and petal primordia. The simulations reproduced the diversity of monocots and eudicots, including snapdragon Antirrhinum majus and its cyc mutant, with respect to organ number, arrangement and initiation patterns, which were dependent on the inhibition strength. These theoretical results suggest that diversity in floral symmetry is primarily regulated by the dorso-ventral inhibitory field and meristem size during developmental evolution.

<i>KEEL LOSS1</i> Regulates Petal Number Along the Floral Dorsoventral Axis in <i>Lotus Japonicus</i> and <i>Pisum Sativum</i>

Although variation of floral organ numbers has been a prevalent phenomenon during the evolution of zygomorphic flowers among many species, most legumes in the subfamily Papilionoideae possess pentamerous corollas. Previous work showed that dorsoventral asymmetrically expressed CYCLOIDEA (CYC)-like genes have been recruited to the establishment of floral zygomorphy. However, the mechanism determining petal number along the dorsoventral axis remains unknown. Here we address this question by a comparative study of floral development between two papilionoid legumes and characterization of two new genetic loci, KEEL LOSS1 (KEL1) from Lotus japonicus and RED BULL1 (REB1) from pea (Pisum sativum L.). kel1 and reb1 mutants show varying degrees of petal absence along the dorsoventral axis, resembling diverse floral morphologies in legumes with non-pentamerous corollas. Ontogenetic analysis shows that the kel1/reb1 phenotype is caused by the defect of early stage common primordia shared between petal and stamen. KEL1 and REB1 are orthologs encoding APETALA2 (AP2) type transcription factors and are both expressed in the ventral or ventral/dorsal region of the floral meristem, consistent with the predominant absence of petals at the corresponding positions in respective mutants. We also find that KEL1 is independent of the CYC pathway, and interacts with PROLIFERATING FLORAL ORGANS (PFO) to control the common primordia indeterminacy by regulating LjWUSCHEL and LjAGAMOUS expression in L. japonicus. We conclude that KEL1/REB1 regulates position-specific floral patterning and confers robust petal numbers along the dorsoventral axis in papilionaceous flowers. While its major role of controlling petal number is largely conserved between L. japonicus and pea, loss of KEL1 in different ontogenetic or genetic background often leads to distinct floral morphologies, suggesting that KEL1 may be a key dorsoventral factor driving the evolution of legume flowers.

Pollination biology in Roepera (Zygophyllaceae): How flower structure and shape influence foraging activity

AbstractThe foraging behavior of bees is a complex phenomenon that depends on numerous physical features of flowers. Of particular importance are accessibility of floral rewards, floral proportions, symmetry and orientation. The flowers of Roepera are characterized by the presence of staminal scales (SS), which play an important role in nectar protection. We studied two species of Roepera with different symmetry and flower orientation, which are mainly visited by honeybees (Apis mellifera). We aimed to show how the foraging behavior of honey bees is affected by the function of SS, floral symmetry and orientation. The foraging behavior was documented by video photography. Handling time, access to nectar, percentage of pollen/nectar foraging, percentage of pollen contact and pollen deposition site on the honey bee's body were assessed. The morphometric features of the honey bees and flowers were analyzed. We found that the SS restricted pollinator access to nectar. Our results indicated consistency of visitation patterns in zygomorphic, laterally oriented flowers of R. fuscata versus random patterns in actinomorphic, diversely oriented flowers of R. leptopetala. The relative proportions of SS and proboscis length appear to be crucial for the success of pollinators. The directionality of the honey bees' movement, together with the different positioning of reproductive organs, plays an important role in the accuracy of pollen transfer and pollination efficiency.

Phylogenetic relationships and biogeographic history of the unique Saxifraga sect. Irregulares (Saxifragaceae) from eastern Asia

AbstractSaxifraga L. is the largest genus in Saxifragaceae and a characteristic component of the herbaceous flora of the temperate and alpine mountains in the Northern Hemisphere. Section Irregulares is a small group of 15–20 species, representing one of the early‐diverged lineages in the genus characterized with unique zygomorphic flowers. We used both nuclear internal transcribed spacer and chloroplast DNA regions (psbA‐trnH, trnL‐F, and matK) to reconstruct its species relationships, estimate divergence times, and infer its historical biogeography. Our phylogenetic results corroborate the monophyly of sect. Irregulares and its sister relationship to sect. Heterisia from North America. The section was well resolved into two lineages corresponding to their morphological features and biogeographic distributions. One represents ser. Stoloniferae including S. stolonifera Curtis and S. nipponica Makino with long‐creeping stolons/rhizomes and small petals with spots and the other comprises the remaining taxa (ser. Rufescentes) which lack long‐creeping rhizomes. Spots on leaves (abaxially spotted vs. abaxially without spots) and spots on petals (without spots vs. with spots) are inferred to be phylogenetically informative within ser. Rufescentes. Divergence time estimates and ancestral area analysis suggested a western North American origin of sect. Irregulares with migration into East Asia by way of the Bering land bridge in the Middle Oligocene. The development of drying and desertification belt in the late Miocene could have played an important role in the subsequent restriction and separation of the north and south lineages within eastern Asia.

Effects of invasive Cirsium arvense on pollination in a southern Appalachian floral community vary with spatial scale and floral symmetry

Invasive plants can alter pollination dynamics by disrupting pollinator visitation and pollen transfer dynamics. However, a consensus regarding the direction of their overall effects (competitive vs. facilitative) remains elusive. Here, we evaluate the role of floral traits and spatial scale (community vs. floral neighborhood) in mediating invasive Cirsium arvense effects on resident plant species at multiple stages of the pollination process. C. arvense decreased pollinator visitation rate at the community level only for species with radial floral symmetry. At the floral neighborhood scale, pollinator visitation rate to all resident species was lower in the presence of C. arvense regardless of symmetry. C. arvense altered patterns of conspecific pollen receipt at the floral neighborhood scale, but the direction of the effect varied by plant species. We argue that these scale-dependent effects may be mediated by differences in foraging range and behavior of the main pollinators in the community. C. arvense, however, did not affect the overall reproductive success of resident species at either scale, suggesting that plants at our study sites may not be pollen limited. We further show evidence suggesting that C. arvense may alter the structure of community-level plant–plant interactions via heterospecific pollen transfer by subverting the roles of resident pollen donors within the pollen transfer network. Overall, our results suggest that generalized species (with radial flowers) may be more susceptible to invasive species’ effects than specialized ones (bilateral flowers) and highlight the need to consider scale-dependent effects in order to develop a more predictive understanding of invasive species effects in nature.

Evolutionary Conservation of the Orchid MYB Transcription Factors DIV, RAD, and DRIF.

The MYB transcription factors DIVARICATA (DIV), DIV-and-RAD-Interacting-Factor (DRIF), and the small interfering peptide RADIALIS (RAD) can interact, forming a regulatory module that controls different plant developmental processes. In the snapdragon Antirrhinum majus, this module, together with the TCP transcription factor CYCLOIDEA (CYC), is responsible for the establishment of floral dorsoventral asymmetry. The spatial gene expression pattern of the OitDIV, OitDRIF, and OitRAD homologs of Orchis italica, an orchid with zygomorphic flowers, has suggested a possible conserved role of these genes in bilateral symmetry of the orchid flower. Here, we have identified four DRIF genes of orchids and have reconstructed their genomic organization and evolution. In addition, we found snapdragon transcriptional cis-regulatory elements of DIV and RAD loci generally conserved within the corresponding orchid orthologues. We have tested the biochemical interactions among OitDIV, OitDRIF1, and OitRAD of O. italica, showing that OitDRIF1 can interact both with OitDIV and OitRAD, whereas OitDIV and OitRAD do not directly interact, as in A. majus. The analysis of the quantitative expression profile of these MYB genes revealed that in zygomorphic orchid flowers, the DIV, DRIF1, and RAD transcripts are present at higher levels in the lip than in lateral inner tepals, whereas in peloric orchid flowers they show similar expression levels. These results indicate that MYB transcription factors could have a role in shaping zygomorphy of the orchid flower, potentially enriching the underlying orchid developmental code.

Effect of soil moisture regimes in the early flowering stage on inflorescence morphology and medicinal ingredients of Chrysanthemum morifolium Ramat. Cv. ‘Hangju’

The quality of the traditional Chinese medicine (TCM) is closely related to the climatic conditions of its medicinal plant’s habitats. When Chrysanthemum morifolium Ramat. cv. ‘Hangju’ (Hj) is in the period of flower-bud differentiation, the rainy season also prevails in its original places. In this study, we set three groups of soil moisture regimes (SMR) corresponding to 35%–40% (S1), 65%–70% (S2), 95%–100% (S3) of soil water holding capacity (WHC) and we found for the first time that the effects of different SMR on the inflorescence of Hj in the early flowering stage. With the increase of SMR content, both ray florets (RFs) and disc florets (DFs) are inhibited on inflorescence morphology, especially for DFs. In addition, a CYC2c orthologous gene in “Zaoyang” (ZY, a cultivar of Hj) was identified and sequenced, the phylogenetic analysis showed that this CYCLOIDEA gene in Asteraceae shared duplications which were well-conserved and lineage-specific. Expression analysis showed that HjCYC2c was mainly expressed in RFs, and high SMR condition led to up-regulation. At the same time, our study suggests that FLS gene is a strong candidate as a key factor in Hj for further interaction studies between flavonoids biosynthesis and regulatory factors of floral symmetry. The determination results of medicinal ingredients showed that DFs were more quickly to respond to the SMR variation and produced more medicinal ingredients compared with RFs.

Specialization in plant\u2013pollinator networks: insights from local-scale interactions in Glenbow Ranch Provincial Park in Alberta, Canada

BackgroundThe occurrence and frequency of plant–pollinator interactions are acknowledged to be a function of multiple factors, including the spatio-temporal distribution of species. The study of pollination specialization by examining network properties and more recently incorporating predictors of pairwise interactions is emerging as a useful framework, yet integrated datasets combining network structure, habitat disturbance, and phylogenetic information are still scarce.ResultsWe found that plant–pollinator interactions in a grassland ecosystem in the foothills of the Rocky Mountains are not randomly distributed and that high levels of reciprocal specialization are generated by biological constraints, such as floral symmetry, pollinator size and pollinator sociality, because these traits lead to morphological or phenological mismatching between interacting species. We also detected that landscape degradation was associated with differences in the network topology, but the interaction webs still maintained a consistently higher number of reciprocal specialization cases than expected. Evidence for the reciprocal evolutionary dependence in visitors (e.g., related pollinators visiting related plants) were weak in this study system, however we identified key species joining clustered units.ConclusionsOur results indicate that the conserved links with keystone species may provide the foundation for generating local reciprocal specialization. From the general topology of the networks, plant–pollinators interactions in sites with disturbance consisted of generalized nodes connecting modules (i.e., hub and numerous connectors). Vice versa, interactions in less disturbed sites consisted of more specialized and symmetrical connections.

Ovule and seed production patterns in relation to flower size variations in actinomorphic and zygomorphic flower species.

Zygomorphic flower species tend to show lower flower size variation than actinomorphic flower species. Have these differences also brought an association in ovule and seed production that has arisen due to natural selection in these species? Flowers were collected from 29 actinomorphic and 20 zygomorphic flower species, and fruits were collected from 21 actinomorphic and 14 zygomorphic flower species in Miyagi and Aomori prefectures, in Japan. The coefficient of variations (CVs) of flower sizes, mean ovule sizes of flowers, ovule numbers of flowers and mean seed sizes of fruits were calculated. The CV of flower sizes was marginally different between the floral symmetry types; tending to be lower in the zygomorphic flower species than in the actinomorphic flower species. The CVs of mean ovule sizes and ovule numbers of flowers increased with increase in the CV of flower sizes in the actinomorphic flower species but not in the zygomorphic flower species. Mean ovule number of flowers tends to increase with increase in mean flower size in the actinomorphic flower species but not in the zygomorphic flower species. The degrees in variations in ovule size and number of flowers were influenced by the interaction of floral symmetry type and flower size variation, suggesting that floral symmetry also has brought an evolutionary association in ovule production by flowers.

Genetic insights into the modification of the pre-fertilization mechanisms during plant domestication.

Plant domestication is the process of adapting plants to human use by selecting specific traits. The selection process often involves the modification of some components of the plant reproductive mechanisms. Allelic variants of genes associated with flowering time, vernalization, and the circadian clock are responsible for the adaptation of crops, such as rice, maize, barley, wheat, and tomato, to non-native latitudes. Modifications in the plant architecture and branching have been selected for higher yields and easier harvests. These phenotypes are often produced by alterations in the regulation of the transition of shoot apical meristems to inflorescences, and then to floral meristems. Floral homeotic mutants are responsible for popular double-flower phenotypes in Japanese cherries, roses, camellias, and lilies. The rise of peloric flowers in ornamentals such as snapdragon and florists' gloxinia is associated with non-functional alleles that control the relative expansion of lateral and ventral petals. Mechanisms to force outcrossing such as self-incompatibility have been removed in some tree crops cultivars such as almonds and peaches. In this review, we revisit some of these important concepts from the plant domestication perspective, focusing on four topics related to the pre-fertilization mechanisms: flowering time, inflorescence architecture, flower development, and pre-fertilization self-incompatibility mechanisms.

Global geographic patterns of heterospecific pollen receipt help uncover potential ecological and evolutionary impacts across plant communities worldwide

Species interactions are known to be key in driving patterns of biodiversity across the globe. Plant-plant interactions through heterospecific pollen (HP) transfer by their shared pollinators is common and has consequences for plant reproductive success and floral evolution, and thus has the potential to influence global patterns of biodiversity and plant community assembly. The literature on HP transfer is growing and it is therefore timely to review patterns and causes of among-species variation in HP receipt at a global scale, thus uncovering its potential contribution to global patterns of biodiversity. Here we analyzed published data on 245 species distributed across five continents to evaluate latitudinal and altitudinal patterns of HP receipt. We further analyzed the role of floral symmetry and evolutionary history in mediating patterns of HP receipt. Latitude and elevation affected the likelihood and intensity of HP receipt indicating that HP transfer increases in species-rich communities and in areas with high abundance of vertebrate pollinators. Floral symmetry and evolutionary history determined HP load size across plant communities worldwide. Overall, our results suggest that HP receipt may have the potential to contribute to global geographic patterns of plant diversity by imposing strong selective pressures in species-rich areas across the globe.

Zygotritonia atropurpurea (Iridaceae: Crocoideae), a new local endemic species from northern Zambia of this small tropical African genus

A seventh species of the tropical and sub-Saharan African genus Zygotritonia Mildbr. has recently been discovered in northern Zambia. It is distinct in the genus in its tall stature, sometimes exceeding 1.3 m, and multibranched spike bearing numerous relatively small, dark purple flowers. The new species, here described as Z. atropurpurea Goldblatt & J.C.Manning, broadly conforms to other species of the genus in its pleated foliage leaf or leaves, zygomorphic flowers with a large, arching dorsal tepal and smaller upper lateral and lower tepals, the three lowermost directed downward and with spooned tips. Known by both morphological comparison and DNA sequence analysis to be a member of tribe Watsonieae of Crocoideae, Zygotritonia is allied to the Lapeirousia Pourr. group of genera, specifically Savannosiphon Goldblatt & Marais and Cyanixia Goldblatt & J.C.Manning.

Phylogenomic analyses reveal an exceptionally high number of evolutionary shifts in a florally diverse clade of African legumes

Detarioideae is well known for its high diversity of floral traits, including flower symmetry, number of organs, and petal size and morphology. This diversity has been characterized and studied at higher taxonomic levels, but limited analyses have been performed among closely related genera with contrasting floral traits due to the lack of fully resolved phylogenetic relationships. Here, we used four representative transcriptomes to develop an exome capture (target enrichment) bait for the entire subfamily and applied it to the Anthonotha clade using a complete data set (61 specimens) representing all extant floral diversity. Our phylogenetic analyses recovered congruent topologies using ML and Bayesian methods. Anthonotha was recovered as monophyletic contrary to the remaining three genera (Englerodendron, Isomacrolobium and Pseudomacrolobium), which form a monophyletic group sister to Anthonotha. We inferred a total of 35 transitions for the seven floral traits (pertaining to flower symmetry, petals, stamens and staminodes) that we analyzed, suggesting that at least 30% of the species in this group display transitions from the ancestral condition reconstructed for the Anthonotha clade. The main transitions were towards a reduction in the number of organs (petals, stamens and staminodes). Despite the high number of transitions, our analyses indicate that the seven characters are evolving independently in these lineages. Petal morphology is the most labile floral trait with a total of seven independent transitions in number and seven independent transitions to modification in petal types. The diverse petal morphology along the dorsoventral axis of symmetry within the flower is not associated with differences at the micromorphology of petal surface, suggesting that in this group all petals within the flower might possess the same petal identity at the molecular level. Our results provide a solid evolutionary framework for further detailed analyses of the molecular basis of petal identity.

Patterns of Diversity of Floral Symmetry in Angiosperms: A Case Study of the Order Apiales

Floral symmetry is widely known as one of the most important structural traits of reproductive organs in angiosperms. It is tightly related to the shape and arrangement of floral parts, and at the same time, it plays a key role in general appearance (visual gestalt) of a flower, which is especially important for the interactions of zoophilous flowers with their pollinators. The traditional classification of floral symmetry divides nearly all the diversity of angiosperm flowers into actinomorphic and zygomorphic ones. Within this system, which is useful for ecological studies, many variations of symmetry appear to be disregarded. At the same time, the diversity of floral symmetry is underpinned not only by ecological factors, but also by morphogenetic mechanisms and constraints. Sometimes it is not an easy task to uncover the adaptive or developmental significance of a change of the floral symmetry in a particular lineage. Using the asterid order Apiales as a model group, we demonstrate that such changes can correlate with the merism of the entire flower or of its particular whorl, with the relative orientation of gynoecium to the rest of the flower, with the presence of sterile floral elements and other morphological characters. Besides, in some taxa, the shape and symmetry of the flower change in the course of its development, which should be taken in consideration in morphological comparisons and evaluations of synapomorphies in a particular clade. Finally, we show that different results can be obtained due to employment of different approaches: for instance, many flowers that are traditionally described as actinomorphic turn out to be disymmetric, monosymmetric, or asymmetric from a more detailed look. The traditional method of division into actinomorphy and zygomorphy deals with the general appearance of a flower, and mainly considers the shape of the corolla, while the geometrical approach handles the entire three-dimensional structure of the flower, and provides an exact number of its symmetry planes.

Diversity of flowers in basic structure and its underlying molecular mechanisms

As the reproductive structures of angiosperms, flowers show tremendous diversity in morphology and structure. The diversification of flowers in basic structure and other features has greatly contributed to the success of angiosperms. In recent years, thanks to the rise of, as well as the rapid progress in evolutionary developmental biology (evo-devo), our knowledge of the molecular basis for the development and diversification of flowers in basic structure has advanced considerably. In this review, we attempt to summarize the developmental and evolutionary mechanisms underlying the diversification of flowers in terms of types, fusion, arrangement patterns, and number of floral organs, as well as floral symmetry. We suggest that future studies should be focused on the establishment of new model systems, application of advanced approaches and integration of multiple disciplines to elucidate the processes, patterns, causes, and molecular bases of the diversification of flowers in basic structure.

Interactions between WUSCHEL- and CYC2-like Transcription Factors in Regulating the Development of Reproductive Organs in Chrysanthemum morifolium.

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.