Pitfall Flower Development and Organ Identity of Ceropegia sandersonii (Apocynaceae-Asclepiadoideae)

Deceptive Ceropegia pitfall flowers are an outstanding example of synorganized morphological complexity. Floral organs functionally synergise to trap fly-pollinators inside the fused corolla. Successful pollination requires precise positioning of flies headfirst into cavities at the gynostegium. These cavities are formed by the corona, a specialized organ of corolline and/or staminal origin. The interplay of floral organs to achieve pollination is well studied but their evolutionary origin is still unclear. We aimed to obtain more insight in the homology of the corona and therefore investigated floral anatomy, ontogeny, vascularization, and differential MADS-box gene expression in Ceropegia sandersonii using X-ray microtomography, Light and Scanning Electronic Microscopy, and RT-PCR. During 10 defined developmental phases, the corona appears in phase 7 at the base of the stamens and was not found to be vascularized. A floral reference transcriptome was generated and 14 MADS-box gene homologs, representing all major MADS-box gene classes, were identified. B- and C-class gene expression was found in mature coronas. Our results indicate staminal origin of the corona, and we propose a first ABCDE-model for floral organ identity in Ceropegia to lay the foundation for a better understanding of the molecular background of pitfall flower evolution in Apocynaceae.

Silencing of FOREVER YOUNG FLOWER-Like Genes from Phalaenopsis Orchids Promotes Flower Senescence and Abscission

Ectopic expression of FOREVER YOUNG FLOWER (FYF) delays floral senescence and abscission in transgenic Arabidopsis. To analyze the FYF function in Phalaenopsis orchids, two FYF-like genes (PaFYF1/2) were identified. PaFYF1/2 were highly expressed in young Phalaenopsis flowers, and their expression decreased significantly afterward until flower senescence. This pattern was strongly correlated with the process of flower senescence and revealed that PaFYF1/2 function to suppress senescence/abscission during early flower development. Interestingly, in flowers, PaFYF1 was consistently expressed less in petals than in lips/sepals, whereas PaFYF2 was expressed relatively evenly in all flower organs. This difference suggests a regulatory modification of the functions of PaFYF1 and PaFYF2 during Phalaenopsis flower evolution. Delayed flower senescence and abscission, which were unaffected by ethylene treatment, were observed in 35S::PaFYF1/2 and 35S::PaFYF1/2 + SRDX transgenic Arabidopsis plants due to the downregulation of the ethylene signaling and abscission-associated genes EDF1-4, IDA and BOP1/2. These results suggest a possible repressor role for Phalaenopsis PaFYF1/2 in controlling floral senescence/abscission by suppressing ethylene signaling and abscission-associated genes. To further validate the function of PaFYF1/2, PaFYF1/2-VIGS (virus-induced gene silencing) Phalaenopsis were generated and analyzed. Promotion of senescence and abscission was observed in PaFYF1/2-VIGS Phalaenopsis flowers by the upregulation of PeEDF1/2, PeSAG39 and PeBOP1/2 expression, the early occurrence of greening according to their increased chlorophyll content and the reduction in water content in flower organs. Our results support that PaFYF1/2 function as transcriptional repressors to prohibit flower senescence and abscission in Phalaenopsis.

Parallelism in Flower Evolution and Development

Flower evolution is characterized by widespread repetition, with adaptations to pollinator environment evolving in parallel. Recent studies have expanded our understanding of the developmental basis of adaptive floral novelties—petal fusion, bilateral symmetry, heterostyly, and floral dimensions. In this article, we describe patterns of trait evolution and review developmental genetic mechanisms underlying floral novelties. We discuss the diversity of mechanisms for parallel adaptation, the evidence for constraints on these mechanisms, and how constraints help explain observed macroevolutionary patterns. We describe parallel evolution resulting from similarities at multiple hierarchical levels—genetic, developmental, morphological, functional—which indicate general principles in floral evolution, including the central role of hormone signaling. An emerging pattern is mutational bias that may contribute to rapid patterns of parallel evolution, especially if the derived trait can result from simple degenerative mutations. We argue that such mutational bias may be less likely to govern the evolution of novelties patterned by complex developmental pathways.

A unique flower in Miocene amber sheds new light on the evolution of flowers

The evolution of flowers is among the foremost topics in evolutionary science. The question for botanists of how flowers evolved exists mainly due to lack of relevant fossil evidence, especially of well-preserved flowers. Dominican amber has yielded abundant fossils (including those of flowers) and thus opens a unique window on flower evolution. Here we report a unique flower preserved in mid-Miocene Dominican amber, Dinganthus pentamera gen. et sp. nov. The flower is actinomorphic, pentamerous, bisexual flower including two bracts, five tepals, 10 stamens, and gynoecium. The stamens are dorsifixed, filamentous, and latrorse. The gynoecium in the centre comprises three portions, namely, a basal gynophore, multiple ovaries in the middle, and an apical style. Supplementing to the developmental and genetic evidence, the unique morphology of Dinganthus suggests that a flower is a condensed shoot with lateral appendages, a long-held belief in botany.

The maleness of larger angiosperm flowers

Flower biomass varies widely across the angiosperms. Each plant species invests a given amount of biomass to construct its sex organs. A comparative understanding of how this limited resource is partitioned among primary (male and female structures) and secondary (petals and sepals) sexual organs on hermaphrodite species can shed light on general evolutionary processes behind flower evolution. Here, we use allometries relating different flower biomass components across species to test the existence of broad allocation patterns across the angiosperms. Based on a global dataset with flower biomass spanning five orders of magnitude, we show that heavier angiosperm flowers tend to be male-biased and invest strongly in petals to promote pollen export, while lighter flowers tend to be female-biased and invest more in sepals to insure their own seed set. This result demonstrates that larger flowers are not simple carbon copies of small ones, indicating that sexual selection via male–male competition is an important driver of flower biomass evolution and sex allocation strategies across angiosperms.

The neural selection and the emergence of ‘beauty canons’ as signaling codes in co-evolving species

Widespread opinion wants beauty to be pleasant and aimless, this assumption biased Darwin's explanation of sexual selection. Conversely, Wallace hypothesized that showy and symmetric sexual traits correlate with vigor and health and he placed ‘aesthetic’ preferences within the natural selection. The controversy has continued until today. To understand the role of beauty canons in communication, the focus was on the flower-pollinator cooperative system as a model, were flower evolution embodies the natural history of pollinators' preferences.Optimum for a signal requires energy efficiency, high signal-to-noise ratio, and intelligibility. It involves pollinator perception mechanisms that, in turn, induce co-evolutionary feedback on signal traits. In fact, the flowers physical and hedonic properties correlate with the basic perceptual, motivational, emotional, and learning mechanisms of pollinators. It is proposed that pollinator behavior, unmasking a preference, reveals the ability to evaluate an expected benefit. Features such as a relative simplicity, redundancy, and regularity of stimuli facilitate perception and memorization and are essential elements for communication between co-evolving species. They improve signaling to satisfy the need for easy and fast recognition. With these properties, a stimulus is adaptive and rewarding per se and may be an ideal conditioned stimulus in associative learning. Among the most conspicuous signals, pollinators learn to recognize and choose those associated with nectar, thus favoring the evolution of flowers that are not only ‘beautiful’ but also ‘honest’ in reporting a reward. Beauty is an emergent property, and studying communication and perception we may understand the origin of some beauty canons.

Characterization of Two AGL6–Like Genes from a Chinese Endemic Woody Tree, Manglietia patungensis (Magnoliaceae) Provides Insight into Perianth Development and Evolution in Basal Angiosperms

Manglietia patungensis (Magnoliaceae) exhibits radially symmetric flowers with perianth consisting of three separate sepaloid tepals in whorl 1 and six petaloid tepals in the inner two whorls, which shows an obvious difference from flowers of most Magnoliaceae species that contain three uniform petaloid tepals whorls, and make it an excellent model for understanding perianth morphology differentiation during early flower evolution. Here, two AGL6 orthologs, MapaAGL6-1 and MapaAGL6-2, were isolated from M. patungensis. Sequence alignment and phylogenetic analyses grouped both genes into the AGL6 lineage. MapaAGL6-1 is expressed only in the perianth whorls, while MapaAGL6-2 is strongly expressed in the perianth whorls but is lowly expressed in gynoecium. Furthermore, ectopic expression of MapaAGL6-1 results in strong complementation phenotypes in the Arabidopsis ap1-10 flower and production of normal floral organs in four floral whorls only with the petal number reduced in whorl 2, while ectopic expression of MapaAGL6-2 only results in petals partly rescued but failing to terminate carpelloid development in Arabidopsis ap1-10 mutant. In addition, the daughter lines generated from a cross between 35S::MapaAGL6-1 transgenic plants showing strong phenotypes and 35S::MapaAGL6-2 transgenic plants showing phenotypic changes produce normal flowers. Our results suggest that MapaAGL6-1 is a reasonable A-function gene controlling perianth identity in Magnoliaceae, which infers from its expression region and complementation phenotypes in Arabidopsis ap1 mutant, while MapaAGL6-2 is mainly involved in petaloid tepal development. Our data also provide a new clue to uncover the perianth development and early evolution in basal angiosperms.

Honeybees prefer novel insect-pollinated flower shapes over bird-pollinated flower shapes

Plant–pollinator interactions have a fundamental influence on flower evolution. Flower color signals are frequently tuned to the visual capabilities of important pollinators such as either bees or birds, but far less is known about whether flower shape influences the choices of pollinators. We tested European honeybee Apis mellifera preferences using novel achromatic (gray-scale) images of 12 insect-pollinated and 12 bird-pollinated native Australian flowers in Germany; thus, avoiding influences of color, odor, or prior experience. Independent bees were tested with a number of parameterized images specifically designed to assess preferences for size, shape, brightness, or the number of flower-like shapes present in an image. We show that honeybees have a preference for visiting images of insect-pollinated flowers and such a preference is most-likely mediated by holistic information rather than by individual image parameters. Our results indicate angiosperms have evolved flower shapes which influence the choice behavior of important pollinators, and thus suggest spatial achromatic flower properties are an important part of visual signaling for plant–pollinator interactions.