Viola philippica develops chasmogamous (CH) and cleistogamous (CL) flowers under different photoperiods, while Viola cornuta only forms CH flowers. Gene expression variations driven by photoperiods that contribute to the CH–CL transition remain largely unknown. Here, we performed comparative transcriptomics between V. philippica and V. cornuta during flower development and revealed differentially expressed genes (DEGs) between CL and CH flowers. Compared to CL flowers in V. philippica under 16-h daylight conditions, DEGs, including auxin response factor (ARF) genes and floral MADS-box genes, were identified in CH flowers under 12-h daylight conditions; however, their homologous genes in V. cornuta were equivalently expressed in CH flowers across both photoperiods. Genes in the modules most significantly correlated with floral types were DEGs between CH and CL flowers in V. philippica, but they were not altered in V. cornuta CH flowers under either photoperiod. Moreover, genes related to fatty acid, flavonoid, and anthocyanin biosynthesis were upregulated in CH flowers, whereas defense response genes were upregulated in CL flowers. Furthermore, the genes co-expressed with the floral B-class MADS-box gene APETALA3 included ARFs, CRABS CLAW, BIGPETALp, TCP14, and UGT87A, whose homologous genes are involved in nectary and floral organ development, including organ identity, size, and coloration. Thus, the putative coordination of genes involved in defense response and auxin signaling pathways, floral organ identity determination and growth, and fatty acid and flavonol biosynthesis in response to photoperiod might regulate CH–CL development in V. philippica, thus providing new insights into the evolution of dimorphic flower development in Viola.

Angiosperm sexual systems are complex and diverse. Androdioecy, the co‐occurrence of males and hermaphrodites, is rare and considered an intermediate evolutionary state between hermaphrodites and dioecy. However, the evolutionary factors that drive androdioecy remain unclear. Based on studies on Berchemia and Tapiscia life histories, it has been observed that species with long sexual reproductive cycles, in which flowers and fruits develop simultaneously, are typically androdioecious. Conversely, species with short sexual reproductive cycles in which the developmental periods of flowers and fruits do not overlap are typically hermaphroditic.Therefore, we hypothesized that a long reproductive cycle leads to the simultaneous development of flowers and fruits, creating a trade‐off in reproductive resources. This trade‐off ultimately drives evolution from hermaphroditism to androdioecy. To test this hypothesis, we conducted artificial experiments to remove fruits and defoliate representative species of Berchemia and Tapiscia to simulate resource release and short age scenarios, respectively. Differences in the size and biomass of the flower parts between the treatment and control groups were compared. Our findings indicate that the simultaneous development of flowers and fruits leads to a tradeoff in reproductive resources, resulting in reduced investment in flowers. Additionally, limited resources are reallocated between female and male functions in bisexual flowers. Interestingly, when reproductive resources fluctuate, female functions are affected to a greater extent, whereas male functions are more resilient to resource changes. These results indicate that when reproductive resources are traded off, it leads to a bias toward male functions in hermaphrodites, promoting the emergence of male individuals and thus forming an androdioecious sexual system.

Most species in the early divergent angiosperm family Annonaceae are apocarpous, with very diverse gynoecial morphologies. Although several Annonaceae genera with apocarpous flowers are known to possess an extragynoecial compitum (EGC) that enables intercarpellary pollen-tube growth to enhance fertilization success, the occurrence of EGC across the whole family remains obscure. Twenty Annonaceae species from all four subfamilies (Anaxagoreoideae, Ambavioideae, Annonoideae, and Malmeoideae) are examined here, with anatomical evidence revealing the occurrence of stigmatic exudate-mediated suprastylar EGC in all four subfamilies. Histological and ontogenetic studies furthermore indicate that trichomes in Cananga and Drepananthus form a confluent zone that connects adjacent stigmas, providing a structural premise for suprastylar EGC. Infrastylar EGC are reported in the Annonaceae for the first time in several genera, including Artabotrys, Annona, and Miliusa, associated with the opening on the ovary ventral groove and/or basal placentation. In addition to the sister genera Isolona and Monodora that are clearly syncarpous, flowers of the distantly related genus Cyathocalyx with a unicarpellate gynoecium have also been hypothesized to be syncarpous. Evidence of carpel vasculature and primordium development does not support that the solitary Cyathocalyx carpel is originated from carpel fusion, however, although the increased number of ovules renders it functionally similar to syncarpy. Gynoecial features, including the extensive occurrence of EGC and the increased number of ovules per carpel (consonant with reduction to a solitary carpel) in Cyathocalyx, may have evolved to overcome limitations associated with apocarpy and possibly contribute to the reproductive success and diversification of the family.

Despite the importance of gamete production this topic has rarely been investigated in angiosperms using comparative approaches. Here, we investigated pollen and ovule numbers per flower in 73 species and 99 populations of Primula comprising both distylous and homostylous reproductive systems. We investigated whether phylogenetic relationships influenced associations between variation in gamete production, floral traits and elevation, and whether the evolutionary breakdown of distyly to homostyly resulted in parallel changes to gamete production. We used a Bayesian approach facilitated by the MCMCglmm method to model pollen and ovule traits across species and determined whether they exhibited phylogenetic signals. We detected significant positive correlations between pollen number and elevation in both the long‐styled and short‐styled morphs (L‐morph and S‐morph, respectively), whereas ovule number was not influenced by elevation. Pollen production was significantly higher in the L‐morph than in the S‐morph, but there was no significant difference between morphs in ovule number. Pollen volume exhibited a positive correlation with the style length of compatible morphs. The transition from distyly to homostyly was associated with significant decreases in pollen production but not ovule number. Our findings demonstrate the importance of elevation on pollen production, perhaps because of selection to improve pollen‐transfer efficiency in uncertain pollinator environments. In contrast, ovule number variation appears to be more constrained by phylogenetic relationships. Our comparative analyses of a well defined angiosperm lineage highlight the complex interactions between intrinsic and extrinsic factors influencing gamete production in plants and emphasize the importance of considering pollen and ovule data separately.

Gene duplications have contributed to the innovation of morphological traits during plant evolution. An outstanding example is the role of CYCLOIDEA2 (CYC2) gene duplications in the formation of the complex structure of Asteraceae capitula. Previous studies have shown that Chrysanthemum lavandulifolium (Fischer ex Trautv.) Makino and a few other species of the Asteraceae harbor two copies of CYC2e. Here, we identified a lineagespecific CYC2e duplication event at the root of the phylogeny of the tribe Anthemideae by analyzing the evolutionary history of CYC2 genes across the Asteraceae. Although the gene expression patterns of CYC2e1 and CYC2e2 were similar in most of floral organs, CYC2e1 promoted ligule elongation by promoting cell expansion, whereas CYC2e2 showed a weak inhibiting effect on the ray‐floret development, and overexpression of CYC2e2 resulted in the maldevelopment of stamens in disc florets. These results indicated differentiated functions of CYC2e1 and CYC2e2 in the capitulum development of the Anthemideae. Given the previous data that identified CYC2g and CYC2d as main regulators in ligule morphogenesis, we comparatively analyzed 10 Asteraceae genomic data, and identified tandem arrays of CYC2g, CYC2d, and CYC2e. We further investigated the regulatory relationships between CYC2g/CYC2d and CYC2e genes, and found that CYC2g can activate the expression of CYC2e1. The findings of this study elucidate the synergistic roles of CYC2 genes in regulating the formation of the Asteraceae capitula and enhance understanding of the mechanism of duplicated gene retention during plant evolution.

Toona fargesii A. Chev., a versatile tree in the Toona genus of the Meliaceae family, is renowned for its exquisite timber and medicinal properties, offering promising benefits. Due to natural regeneration obstacles and long‐term excessive exploitation, it has been threatened in China. Intriguingly, root sprouting, which may diminish the genetic diversity and hinder population development, dominates the reproductive pattern of T. fargesii in the wild. However, the lack of complete genome information has hampered basic studies on the regeneration, classification, evolution and conservation of this species. Here, we report the genome of T. fargesii, which was sequenced using the PacBio platform and assembled into a high‐quality genome with a total size of 535.24 Mb. Of this, 97.93% of the assembled contigs were anchored onto 28 pseudochromosomes, achieving a chromosomelevel genome. The long terminal repeat assembly index score was 21.34, and the consensus quality value was 39.90%, indicating the accuracy and completeness of the genome. Comparative genome analysis suggested that a recent whole genome duplication event occurred between 22.1 and 50.1 Mya in the Toona genus, with the divergence time between T. fargesii and its relative T. sinensis estimated at approximately ~16.7 Mya. Additionally, 13 TfARR genes, which play integral roles in root sprouting by mediating cytokinin signaling, underwent rapid gene expansion and showed significant enrichment in the plant hormone signal transduction pathway. Furthermore, transcriptomic analysis demonstrated that differentially expressed genes between root sprouts and nonroot sprouts were significantly enriched in the zeatin biosynthesis pathway, indicating that cytokinin regulation is involved in root sprouting development. Collectively, the findings provide valuable genomic resources for the Toona genus and genetic insights into the mechanisms of root sprouting in T. fargesii.

Members of the genus Juglans (walnuts, family Juglandaceae) are typical Tertiary relict tree species. They are widely distributed in China and are well known for their edible nuts and high‐quality wood. The genetic structure and historical gene flow patterns of Chinese Juglans still need to be resolved. Here, we collected samples from a total of 2242 trees from 142 populations of three walnut species (Juglans regia L., Juglans sigillata Dode, and Juglans mandshurica Maxim.) and a hybrid taxon Juglans × hopeiensis. These samples were distributed throughout China. We analyzed population genetic structure, interspecific relationships, lineage differentiation, and speciation using 21 EST‐SSR genetic markers. All populations of J. regia and J. sigillata clustered into one lineage, corresponding to the Juglans sect. Dioscaryon, and all J. mandshurica and Juglans × hopeiensis populations (section Cardiocaryon) clustered into a second lineage. For J. mandshurica, there was an obvious north-south genetic cline. Interestingly, except for seven populations of the hybrid Juglans × hopeiensis, there was almost no gene flow between the populations of section Juglans/Dioscaryon and section Cardiocaryon. A DIYABC analysis showed that J. regia and J. mandshurica differentiated during the Oligocene. Juglans sigillata originated during the Pliocene to the Pleistocene. Subsequently, during the Middle Pleistocene, J. regia and J. mandshurica hybridized within a narrow zone to produce Juglans × hopeiensis. These results demonstrate the impact of repeated turbulent climate changes in the Quaternary on the evolutionary history of Tertiary relict plants.

Polyploids are common in Camellia sect. Paracamellia, which contain many important oil crop species. However, their complex evolutionary history is largely unclear. In this study, 22 transcriptomes and 19 plastomes of related species of Camellia were sequenced and assembled, providing the most completed taxa sampling of Camellia sect. Oleifera and C. sect. Paracamellia. Phylogenetic trees were reconstructed with predicted single-copy nuclear genes and plastomes. Phylogenetic trees with nuclear genes demonstrated that C. sect. Oleifera should be merged into C. sect. Paracamellia. Cytonuclear discordance and network analyses suggested hybridizations among polyploid species and relatives. The divergence of major clades in C. sect. Paracamellia was dated to be during the middle to late Miocene from the ancestral Lingnan region, and a rapid diversification during the Quaternary was found, probably through hybridization and polyploidization. The tetraploid Camellia meiocarpa Hu may have originated from hybridization between closely related diploid species. The hexaploid Camellia oleifera C. Abel probably originated from hybridization between closely related diploid and tetraploid (e.g., C. meiocarpa) species. The octoploid Camellia vietnamensis T. C. Huang ex Hu could have originated from hybridization between hexaploid C. oleifera and the closely related diploid species. Hybridization and polyploidization played an important role in generating the rich variation of important fruit traits, especially increased fruit size in polyploid species.