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Flavonoids, carotenoids, betalains, and chlorophylls are the plant pigments responsible for floral color. Anthocyanins, a class of flavonoids, are largely responsible for the red, purple, pink, and blue colors. R2R3-MYB genes belonging to subgroup 6 (SG6) are the upstream regulatory factors of the anthocyanin biosynthetic pathway. The canonical members of these genes in Arabidopsis include AtMYB75 , AtMYB90 , AtMYB113 , and AtMYB114 . The Aristolochiaceae is an angiosperm lineage with diverse floral groundplans and perianth colors. Saruma henryi exhibits a biseriate perianth with green sepals and yellow petals. All other genera have sepals only, with colors ranging from green (in Lactoris ) to a plethora of yellow to red and purple mixtures. Here, we isolated and reconstructed the SG6 R2R3-MYB gene lineage evolution in angiosperms with sampling emphasis in Aristolochiaceae. We found numerous species-specific duplications of this gene lineage in core eudicots and local duplications in Aristolochiaceae for Saruma and Asarum . Expression of SG6 R2R3-MYB genes examined in different developmental stages and plant organs of four Aristolochiaceae species, largely overlaps with red and purple pigments, suggesting a role in anthocyanin and flavonoid synthesis and accumulation. A directed RNA-seq analysis corroborated our RT-PCR analyses, by showing that these structural enzymes activate during perianth development in Aristolochia fimbriata and that the regulatory genes are expressed in correlation with color phenotype. Finally, the reconstruction of the flavonoid and anthocyanin metabolic pathways using predicted peptides from transcriptomic data show that all pivotal enzymes are present in the analyzed species. We conclude that the regulatory genes as well as the biosynthetic pathway are largely conserved across angiosperms. In addition, the Aristolochiaceae emerges as a remarkable group to study the genetic regulatory network for floral color, as their members exhibit an outstanding floral diversity with elaborate color patterns and the genetic complement for SG6 R2R3-MYB genes is simpler than in core eudicot model species.

• Hydrangea sepals exhibit a wide range of colors, from red, through purple, to blue; the purple color is a color mosaic. However, all of these colors are derived from the same components: simple anthocyanins, 3-O-glycosyldelphinidins, three co-pigment components, acylquinic acids and aluminum ions (Al3+). We show the color mosaic is a result of graded differences in intravacuolar factors. • In order to clarify the mechanisms of mosaic color, we performed single-cell analyses of vacuolar pH, and anthocyanin, co-pigment and Al3+ content. From the sepals, a protoplast mixture of various colors was obtained. The cell color was evaluated by microspectrophotometry and vacuolar pH then was recorded by using a pH microelectrode. The organic and Al3+ contents were quantified by micro-HPLC. • We found that the bluer the cell, the greater the ratio of 5-O-acylquinic acids and Al3+ to anthocyanins. Furthermore, reproducing experiments were conducted by mixing the components under various pH condition; all the colors could be reproduced in the various mixing conditions. • Based on the above, we provide experimental evidence for cell color variation in hydrangea. Our study demonstrates the expression of phenotypic differences without any direct genomic control.

Key messageArabidopsis ETHYLENE RESPONSE FACTOR12 (ERF12), the rice MULTIFLORET SPIKELET1 orthologue pleiotropically affects meristem identity, floral phyllotaxy and organ initiation and is conserved among angiosperms.Reproductive development necessitates the coordinated regulation of meristem identity and maturation and lateral organ initiation via positive and negative regulators and network integrators. We have identified ETHYLENE RESPONSE FACTOR12 (ERF12) as the Arabidopsis orthologue of MULTIFLORET SPIKELET1 (MFS1) in rice. Loss of ERF12 function pleiotropically affects reproductive development, including defective floral phyllotaxy and increased floral organ merosity, especially supernumerary sepals, at incomplete penetrance in the first-formed flowers. Wildtype floral organ number in early formed flowers is labile, demonstrating that floral meristem maturation involves the stabilisation of positional information for organogenesis, as well as appropriate identity. A subset of erf12 phenotypes partly defines a narrow developmental time window, suggesting that ERF12 functions heterochronically to fine-tune stochastic variation in wild type floral number and similar to MFS1, promotes meristem identity. ERF12 expression encircles incipient floral primordia in the inflorescence meristem periphery and is strong throughout the floral meristem and intersepal regions. ERF12 is a putative transcriptional repressor and genetically opposes the function of its relatives DORNRÖSCHEN, DORNRÖSCHEN-LIKE and PUCHI and converges with the APETALA2 pathway. Phylogenetic analysis suggests that ERF12 is conserved among all eudicots and appeared in angiosperm evolution concomitant with the generation of floral diversity.

Heptacodium miconioides Rehd., commonly known as “seven-son flower,” is an ornamental species with a beautiful flower pattern and persistent sepals. Its sepals are of horticultural value, turning bright red and elongating in the autumn; however, the molecular mechanisms that cause sepal color change remain unclear. We analyzed the dynamic changes in anthocyanin composition in the sepal of H. miconioides at four developmental stages (S1-S4). A total of 41 anthocyanins were detected and classified into 7 major anthocyanin aglycones. High levels of the pigments cyanidin-3,5-O-diglucoside, cyanidin-3-O-galactoside, cyanidin-3-O-glucoside, and pelargonidin-3-O-glucoside were responsible for sepal reddening. Transcriptome analysis revealed 15 differentially expressed genes involved in anthocyanin biosynthesis that were detected between 2 developmental stages. Of these, the high expression of HmANS was considered critical structural gene related to anthocyanin biosynthesis pathway in the sepal through co-expression analysis with anthocyanin content. In addition, a transcription factor (TF)-metabolite correlation analysis revealed that three HmMYB, two HmbHLH, two HmWRKY, and two HmNAC TFs exhibited a strong positive role in the regulation of the anthocyanin structural genes (Pearson’s correlation coefficient > 0.90). Luciferase activity assay showed that HmMYB114, HmbHLH130, HmWRKY6, and HmNAC1 could activate the promoters of HmCHS4 and HmDFR1 genes in vitro . These findings increase our understanding of anthocyanin metabolism in the sepal of H. miconioides and provide a guide for studies involving sepal color conversion and regulation.

We previously found that B and AGL6 proteins form L (OAP3-2/OAGL6-2/OPI) and SP (OAP3-1/OAGL6-1/OPI) complexes to determine lip/sepal/petal identities in orchids. Here, we show that the functional L’ (OAP3-1/OAGL6-2/OPI) and SP’ (OAP3-2/OAGL6-1/OPI) complexes likely exist and AP3 / PI/AGL6 genes have acquired additional functions during evolution. We demonstrate that the presumed L’ complex changes the structure of the lower lateral sepals and helps the lips fit properly in the center of the flower. In addition, we find that OAP3-1/OAGL6-1/OPI in SP along with presumed SP’ complexes regulate anthocyanin accumulation and pigmentation, whereas presumed L’ along with OAP3-2/OAGL6-2/OPI in L complexes promotes red spot formation in the perianth. Furthermore, the B functional proteins OAP3-1/OPI and OAGL6-1 in the SP complex could function separately to suppress sepal/petal senescence and promote pedicel abscission, respectively. These findings expand the current knowledge behind the multifunctional evolution of the B and AGL6 genes in plants. B class AP3/PI and AGL6-like MADS proteins determine lips and sepals/petals identities in orchids. Here, the authors characterize the extended function of OAP3/OPI/OAGL6 in regulating the specific structure of the lateral sepals, pigmentation/senescence of the perianth and abscission of the pedicel.

APETALA2 (AP2) is best known for its function in the outer two floral whorls, where it specifies the identities of sepals and petals by restricting the expression of AGAMOUS (AG) to the inner two whorls in Arabidopsis thaliana. Here, we describe a role of AP2 in promoting the maintenance of floral stem cell fate, not by repressing AG transcription, but by antagonizing AG activity in the center of the flower. We performed a genetic screen with ag-10 plants, which exhibit a weak floral determinacy defect, and isolated a mutant with a strong floral determinacy defect. This mutant was found to harbor another mutation in AG and was named ag-11. We performed a genetic screen in the ag-11 background to isolate mutations that suppress the floral determinacy defect. Two suppressor mutants were found to harbor mutations in AP2. While AG is known to shut down the expression of the stem cell maintenance gene WUSCHEL (WUS) to terminate floral stem cell fate, AP2 promotes the expression of WUS. AP2 does not repress the transcription of AG in the inner two whorls, but instead counteracts AG activity.

Eight new species of Hymenochilus from Australia are described and illustrated. Six of the new species have affinities to (and are here compared with) H. cycnocephalus: H. anemophilus, H. calcicola, H. cymbellus, H. longipes, H. nemoralis and H. pachylus. Hymenochilus anemophilus is shorter (2–8 cm), with dark green strongly veined rosette leaves, crowded dark green flowers, oblong to obovate labellum lamina and, a broader blunter beak on the labellum appendage; H. calcicola is shorter (3–12 cm), crowded green flowers with prominent dark green stripes, ovate petals with a strongly developed basal flange on the anterior side and narrow elliptic labellum lamina with a broad pointed beak on the labellum appendage; H. cymbellus differs by its sparser basal rosette with narrower rosette leaves, thinner scapes, flowers with distinct darker green stripes and shallowly saccate lateral sepals that narrow inwards to a distinctly pointed apex; H. longipes differs by its thin-textured rosette leaves, widely spaced darker green flowers with darker green veins, elliptic-obovate labellum lamina and a longer labellum basal appendage which protrudes prominently from the flower in side view; H. nemoralis has longer rosette leaves, thicker scape  and darker green flowers with prominent narrow dark green stripes; H. pachylus has thicker rosette leaves, taller, thicker scape, flowers prominently striped, elliptic-obovate labellum lamina and labellum appendage with a short thick beak. Two of the new species have affinities (and are here compared) with H. muticus: H. pagophilus and H. pisinnus. Hymenochilus pagophilus differs by its moderately crowded to crowded flowers, broader, shinier flowers and rectangular-obovate labellum and H. pisinnus differs by its smaller rosette with smaller, narrower leaves, thinner scape, smaller flowers, that are often on long pedicels, shorter, shallowly saccate lateral sepals, smaller rhomboid petals and, smaller obovate labellum. In addition, Hymenochilus cycnocephalus and H. muticus, are characterised in the strict sense with full descriptions, distribution, and habitat. 

A new species, Impatiens ngariensis (Balsaminaceae) from western Xizang, China, is described in this study. Morphological and phylogenetic evidence supports its taxonomic placement within I. sect. Racemosae . Phylogenetic analyses indicate that I. ngariensis forms a clade with I. thomsonii , I. bomiensis , I. fragicolor , I. edgeworthii and I. glandulifera . Morphologically, the new species closely resembles I. sulcata in petiole with 2 basal glands, subcorymbose-racemose inflorescences, lateral sepals ovate, dorsal petal suborbicular and capsule linear, but can be distinguished by several key characteristics, including a suborbicular dorsal petal with a cordate base and a shorter spur.

Cymbidium ensifolium is one of the national orchids in China, which has high ornamental value with changeable flower colors. To understand the formation mechanism of different flower colors of C. ensifolium, this research conducted transcriptome and metabolome analyses on four different colored sepals of C. ensifolium. Metabolome analysis detected 204 flavonoid metabolites, including 17 polyphenols, 27 anthocyanins, 75 flavones, 34 flavonols, 25 flavonoids, 18 flavanones, and 8 isoflavones. Among them, purple-red and red sepals contain a lot of anthocyanins, including cyanidin, pelargonin, and paeoniflorin, while yellow-green and white sepals have less anthocyanins detected, and their metabolites are mainly flavonols, flavanones and flavonoids. Transcriptome sequencing analysis showed that the expression levels of the anthocyanin biosynthetic enzyme genes in red and purple-red sepals were significantly higher than those in white and yellow-green sepals of C. ensifolium. The experimental results showed that CeF3′H2, CeDFR, CeANS, CeF3H and CeUFGT1 may be the key genes involved in anthocyanin production in C. ensifolium sepals, and CeMYB104 has been proved to play an important role in the flower color formation of C. ensifolium. The results of transformation showed that the CeMYB104 is involved in the synthesis of anthocyanins and can form a purple-red color in the white perianth of Phalaenopsis. These findings provide a theoretical reference to understand the formation mechanism of flower color in C. ensifolium.Key messageThis study identified the differential metabolites and differential genes among different color sepals, determined the key regulatory genes, and constructed a regulatory network for the flower color formation of Cymbidium ensifolium.

Orchids are renowned for their intricate floral structures, where sepals and petals contribute significantly to ornamental value and pollinator attraction. In Section , the distinctive curvature of these floral organs enhances both aesthetic appeal and pollination efficiency. However, the molecular and cellular mechanisms underlying this trait remain poorly understood. Morphological characteristics of five hybrids were analyzed, with a particular focus on hybrid H5, which exhibits pronounced sepal curling. Full-length transcriptomic sequencing was employed to assemble a reference transcriptome, while RNA-seq identified differentially expressed genes (DEGs) between sepals and petals. Gene ontology and pathway enrichment analyses were conducted to uncover biological processes associated with sepal curvature. Cytological microscopy was used to examine cell size and number, and quantitative real-time PCR (qRT-PCR) was performed to validate transcriptomic findings. The reference transcriptome contained 94,258 non-redundant transcripts, and RNA-seq identified 821 DEGs between sepals and petals, with 72.8% of these upregulated in sepals. Enrichment analysis revealed the significant involvement of DEGs in cytokinesis, cytoskeletal organization, and energy metabolism. Notably, myosin II filament organization was implicated in generating the mechanical forces responsible for curling, while metabolic pathways provided the energy necessary for these developmental processes. Cytological observations showed that the upper cell layers of the sepal were smaller and more numerous than the lower layers, indicating that differential cell growth contributes to sepal curvature. qRT-PCR analysis validated the differential expression of selected genes, supporting the transcriptomic findings. The interplay of cellular mechanics, cytoskeletal dynamics, and metabolic regulation is crucial in shaping sepal morphology. Future studies involving gene knockdown or overexpression experiments are recommended to validate the roles of specific genes in processes such as actin organization and myosin activity. Such work would provide deeper insights into the contributions of cytoskeletal dynamics and mechanical force generation to sepal morphogenesis.