Search

Understanding hybridization and introgression between natural plant populations can give important insights into the origins of cultivated species. Recent studies suggest differences in ploidy may not create such strong reproductive barriers as once thought, and thus studies into cultivated origins should examine all co-occurring taxa, including those with contrasting ploidy levels. Here, we characterized hybridization between Chrysanthemum indicum, Chrysanthemum vestitum and Chrysanthemum vestitum var. latifolium, the most important wild species involved in the origins of cultivated chrysanthemums. We analysed population structure of 317 Chrysanthemum accessions based on 13 microsatellite markers and sequenced chloroplast trnL-trnF for a subset of 103 Chrysanthemum accessions. We identified three distinct genetic clusters, corresponding to the three taxa. We detected 20 hybrids between species of different ploidy levels, of which 19 were between C. indicum (4x) and C. vestitum (6x) and one was between C. indicum and C. vestitum var. latifolium (6x). Fourteen hybrids between C. indicum and C. vestitum were from one of the five study sites. Chrysanthemum vestitum and C. vestitum var. latifolium share only one chloroplast haplotype. The substantially different number of hybrids between hybridizing species was likely due to different levels of reproductive isolation coupled with environmental selection against hybrids. In addition, human activities may play a role in the different patterns of hybridization among populations.

Most Rosaceae flowers are pentamerous and have petals, but subtribe Sanguisorbinae have small tetramerous (or trimerous) flowers without petals, and their floral morphology and morphogenesis remain poorly known. We investigated the floral development of three Sanguisorba spp. using scanning electron microscopy to clarify the relationship between floral development and mature flowers, with emphasis on floral constraints affecting reduction of petals, diversity of androecial patterns and the development of the gynoecium and ovule, and to clarify the pollination mechanisms in the absence of petals. All species have botryoids, with numerous flowers initiated acropetally, and the maturation of flowers follows different directions. All flowers are tetramerous and sepals emerge pairwise. No petal primordia appear. Antesepalous stamens are initiated in a pairwise manner; only S. hakusanensis has a second whorl. The ovary appears inferior by development of a cystiform hypanthium. There is only one ovule with a single integument by reduction of the inner. The space occupied by the larger sepals forces the stamens to develop sequentially. Our data demonstrate that petals are basically absent and that there is a possible shift of pollination mechanisms in Sanguisorba among the mainly wind-pollinated Sanguisorbinae, as both anemophilous and entomophilous characters are found in this genus.

INTRODUCTION Chrysanthemums are traditional flowers that originated in China and have high ornamental, economic and medicinal value. They are widely used as herbal remedies and consumed as food or beverages in folk medicine. However, little is known about their metabolic composition. OBJECTIVES The aims of this work were to determine the metabolic composition of and natural variation among different species of Chrysanthemum and to explore new potential resources for drug discovery and sustainable utilisation of wild Chrysanthemum. METHODS The metabolomes of Chrysanthemum mongolicum (Ling) Tzvel. and Chrysanthemum rhombifolium H. Ohashi & Yonek. were compared using a widely targeted metabolomics approach based on liquid chromatography-tandem mass spectrometry (LC-MS/MS). RESULTS In total, 477 metabolites were identified, of which 72 showed significant differences in expression between C. mongolicum and C. rhombifolium, mainly in flavonoids, organic acids and nucleotides. The flavone and flavonol biosynthesis pathway showed significant enrichment among the differentially expressed metabolites. The contents of genkwanin, trigonelline, diosmin, narcissoside, 3,4-dihydroxyphenylacetic acid, linarin, N',N'-p-coumarin, C-hexosyl-tricetin O-pentoside, chrysoeriol, acacetin and kaempferol-3-O-gentiobioside were significantly different between the two species and represent potential biomarkers. CONCLUSION The types of flavonoid-related metabolites in the flavonoid biosynthesis pathway differed between C. mongolicum and C. rhombifolium. The mechanisms underlying the unique adaptations of these two species to their environments may involve variations in the composition and abundance of flavonoids, organic acids, and nucleotides. These methods are promising to identify functional compounds in Chrysanthemum species and can provide potential resources for drug discovery and the sustainable utilisation of Chrysanthemum plants.

iArgyranthemum frutescens/i, which belongs to the Anthemideae (Asteraceae), is widely cultivated as an ornamental plant. In this study, the complete chloroplast genome ofiA. frutescens/iwas obtained based on the sequences generated by Illumina HiSeq. The chloroplast genome ofiA. frutescens/iwas 149,626 base pairs (bp) in length, containing a pair of inverted repeats (IR, 24,510 bp) regions separated by a small single-copy (SSC, 18,352 bp) sequence and a large single-copy (LSC, 82,254 bp) sequence. The genome contained 132 genes, consisting of 85 coding DNA sequences, 37 tRNA genes, and 8 rRNA genes, with nineteen genes duplicated in the IR region. A comparison chloroplast genome analysis among ten species from the tribe of Anthemideae revealed that the chloroplast genome size varied, but the genome structure, gene content, and oligonucleotide repeats were highly conserved. Highly divergent regions, e.g.,iycf1/i,itrnK/i-ipsbK/i,ipetN/i-ipsbM/iintronic, were detected. Phylogenetic analysis supportediArgyranthemum/ias a separate genus. The findings of this study will be helpful in the exploration of the phylogenetic relationships of the tribe of Anthemideae and contribute to the breeding improvement ofiA. frutescens/i.

Understanding hybridization and introgression between natural plant populations can give important insights into the origins of cultivated species. Recent studies suggest differences in ploidy may not create such strong reproductive barriers as once thought, and thus studies into cultivated origins should examine all co-occurring taxa, including those with contrasting ploidy levels. Here, we characterized hybridization between Chrysanthemum indicum, Chrysanthemum vestitum and Chrysanthemum vestitum var. latifolium, the most important wild species involved in the origins of cultivated chrysanthemums. We analysed population structure of 317 Chrysanthemum accessions based on 13 microsatellite markers and sequenced chloroplast trnL-trnF for a subset of 103 Chrysanthemum accessions. We identified three distinct genetic clusters, corresponding to the three taxa. We detected 20 hybrids between species of different ploidy levels, of which 19 were between C. indicum (4x) and C. vestitum (6x) and one was between C. indicum and C. vestitum var. latifolium (6x). Fourteen hybrids between C. indicum and C. vestitum were from one of the five study sites. Chrysanthemum vestitum and C. vestitum var. latifolium share only one chloroplast haplotype. The substantially different number of hybrids between hybridizing species was likely due to different levels of reproductive isolation coupled with environmental selection against hybrids. In addition, human activities may play a role in the different patterns of hybridization among populations.

Chrysanthemum rhombifolium (Ling et C. Shih), an endemic plant that is extremely well-adapted to harsh environments. However, little is known about its molecular biology of the plant's resistant traits against stress, or even its molecular biology of overall plant. To investigate the molecular biology of C. rhombifolium and mechanism of stress adaptation, we performed transcriptome sequencing of its leaves using an Illumina platform. A total of 130,891 unigenes were obtained, and 97,496 (~74.5%) unigenes were annotated in the public protein database. The similarity search indicated that 40,878 and 74,084 unigenes showed significant similarities to known proteins from NCBI non-redundant and Swissprot protein databases, respectively. Of these, 56,213 and 42,005 unigenes were assigned to the Gene Ontology (GO) database and Cluster of Orthologous Groups (COG), respectively, and 38,918 unigenes were mapped into five main categories, including 18 KEGG pathways. Metabolism was the largest category (23,128, 59.4%) among the main KEGG categories, suggesting active metabolic processes in C. rhombifolium. About 2,459 unigenes were annotated to have a role in defense mechanism or stress tolerance. Transcriptome analysis of C. rhombifolium revealed the presence of 12,925 microsatellites in 10,524 unigenes and mono, trip, and dinucleotides having higher polymorphism rates. The phylogenetic analysis based on GME gene among related species confirmed the reliability of the transcriptomic data. This work is the first genetic study of C. rhombifolium as a new plant resource of stress-tolerant genes. This large number of transcriptome sequences enabled us to comprehensively understand the basic genetics of C. rhombifolium and discover novel genes that will be helpful in the molecular improvement of chrysanthemums.

Leucanthemella linearis is an important marsh plant. The complete chloroplast genome sequence of L. linearis was obtained using next generation sequencing. It was 15,1401 bp in length, including a pair of inverted repeat (IR, 24,941 bp) regions separated by a small single copy (SSC, 18,392 bp) sequence and a large single copy (LSC, 83,127 bp) sequence. The cp genome contained 140 genes, consisting of 96 protein-coding genes, 8 rRNA genes and 36 tRNA genes. Twenty-two genes were present in the IR region. Thirty-four SSR sites were detected in the cp genome. Phylogenetic analysis with the reported chloroplast genomes revealed that L. linearis is most closely related to Tribe Heliantheae species. This new data will help to understand the phylogenetic position and biology of the Leucanthemella.

Phylogenetic studies have shown that most clades in Prunus are well-supported by the flower structure, but most taxa in the racemose group have not yet been re-evaluated and could contribute to the understanding of the systematic relationships of the subgenera. We examined the inflorescence and flower development in Prunus laurocerasus L. (subgenus Laurocerasus) and P. serotina Ehrh. (subgenus Padus I) using scanning electron microscopy. Our results indicate that they share several floral development characters but differ in the following aspects: (i) all of their flowers are fully developed and each flower is enclosed by a bract and two bracteoles, which later stop development (vs. the terminal flower degenerates and only a single bract subtends each flower); (ii) the style protrudes from the floral bud (vs. the style is crooked and below the anthers); (iii) the outer integument initiates close to the inner one (vs. in the middle of the ovule); and (iv) an obturator appears after initiation of the two integuments (vs. simultaneously with the inner integument). Although our results are preliminary, differences in floral developmental characters support the different origins of Prunus subgenera Laurocerasus and Padus as based on molecular phylogenetic studies.

Background and aims Camptotheca is endemic to China and there are limited data about the breeding system and morphogenesis of the flowers. Camptotheca is thought to be related to Nyssa and Davidia in Nyssaceae, which has sometimes been included in Cornaceae. However, molecular phylogenetic studies confirmed the inclusion of Camptotheca in Nyssaceae and its exclusion from Cornaceae. The aim of this study was to reveal developmental features of the inflorescence and flowers in Camptotheca to compare with related taxa in Cornales. Methods Inflorescences and flowers of Camptotheca acuminata at all developmental stages were collected and studied with a scanning electron microscope and stereo microscope. Key results Camptotheca has botryoids which are composed of several capitate floral units (FUs) that are initiated acropetally. On each FU, flowers are grouped in dyads that are initiated acropetally. All floral organs are initiated centripetally. Calyx lobes are restricted to five teeth. The hypanthium, with five toothed calyx lobes, is adnate to the ovary. The five petals are free and valvate. Ten stamens are inserted in two whorls around the central depression, in which the style is immersed. Three carpels are initiated independently but the ovary is syncarpous and unilocular. The ovule is unitegmic and heterotropous. Inflorescences are functionally andromonoecious varying with the position of the FUs on the inflorescence system. Flowers on the upper FU often have robust styles and fully developed ovules. Flowers on the lower FU have undeveloped styles and aborted ovules, and the flowers on the middle FU are transitional. Conclusions Camptotheca possesses several traits that unify it with Nyssa, Mastixia and Diplopanax. Inflorescence and floral characters support a close relationship with Nyssaceae and Mastixiaceae but a distant relationship with Cornus. Our results corroborate molecular inferences and support a separate family Nyssaceae.

Most multi-objective immune algorithms (MOIAs) adopt clonal selection to speed up convergence, as this operator only clones the best individuals during the search process. However, this approach somehow deteriorates the population diversity, which may cause a MOIA to be trapped in a local optimum and could also lead to premature convergence when tackling some complicated multi-objective optimization problems (MOPs). In order to overcome this problem, an adaptive immune-inspired multi-objective algorithm (AIMA) is presented in this paper, in which multiple differential evolution (DE) strategies having distinct advantages are embedded into a conventional MOIA. Our proposed approach strengthens the exploration capabilities of a MOIA while also improving its population diversity. At each generation, based on the current search stage, an adaptive selection method is designed to choose an appropriate DE strategy for evolution. The core idea is to effectively combine the advantages of three DE strategies when solving different MOPs. A number of comparative experiments are conducted on the well-known and frequently-used WFG and DTLZ test problems. Our experimental results validate the superiority of our proposed AIMA, as it performs better than some state-of-the-art multi-objective optimization algorithms and some state-of-the-art MOIAs.

Recently, evolutionary algorithm based on decomposition (MOEA/D) has been found to be very effective and efficient for solving complicated multiobjective optimization problems (MOPs). However, the selected differential evolution (DE) strategies and their parameter settings impact a lot on the performance of MOEA/D when tackling various kinds of MOPs. Therefore, in this paper, a novel adaptive control strategy is designed for a recently proposed MOEA/D with stable matching model, in which multiple DE strategies coupled with the parameter settings are adaptively conducted at different evolutionary stages and thus their advantages can be combined to further enhance the performance. By exploiting the historically successful experience, an execution probability is learned for each DE strategy to perform adaptive adjustment on the candidate solutions. The proposed adaptive strategies on operator selection and parameter settings are aimed at improving both of the convergence speed and population diversity, which are validated by our numerous experiments. When compared with several variants of MOEA/D such as MOEA/D, MOEA/D-DE, MOEA/D-DE+PSO, ENS-MOEA/D, MOEA/D-FRRMAB and MOEA/D-STM, our algorithm performs better on most of test problems. A novel adaptive control strategy is designed for MOEA/D.The parameter settings are adaptively adjusted at different evolutionary stages.The advantages of four classical differential evolution strategies are combined.Our algorithm gives better performance in solving complicated UF test problems.

Flowering is important for plant propagation and survival, and it is also closely related to human life. Identifying the molecular mechanisms underlying flower development is essential for plant improvement and breeding. Flower development is a complex physiological process that is regulated by multiple genes. LFY genes play important roles in the floral meristem transition and act as crucial integrators in regulating the floral gene network. Argyranthemum frutescens is an ornamental species cultivated for floral displays, yet little is known about molecular mechanisms driving its flower development. In this study, the LEAFY gene homologue, AfLFY, was identified and cloned from A. frutescens, and its role and expression patterns were characterized. Two distinct copies of AfLFY were found in the A. frutescens genome and both sequences contained a 1248 bp open reading frame that encoded 415 amino acids. The putative protein sequences have a typical LFY family domain. In addition, AfLFY was expressed at the highest levels in young leaves of the vegetative stage and in the shoot apical bud meristem of the reproductive stage. Phylogenetic analysis showed that AfLFY was most closely related to DFL from Chrysanthemum lavandulifolium. Subcellular localization studies revealed that AfLFY localized to the nucleus. Heterologous expression of AfLFY in transgenic tobacco plants shortened its period of vegetative growth, converted the lateral meristems into terminal flowers and promoted precocious flowering. In addition, transgenic plants exhibited obvious morphological changes in leaf shape. qRT-PCR analysis indicated that the expression levels genes related to flowering, FT, SOC1, and AP1 were significantly upregulated in AfLFY transgenic plants. Our findings suggested that the AfLFY gene plays a vital role in promoting flowering and leaf development in A. frutescens. These results laid a foundation for us to understand the mechanism of AfLFY in regulation flowering, and the results will be helpful in improving A. frutescens through molecular breeding.

The chrysanthemums (Chrysanthemum × morifolium Ramat.) are a well-known group of traditional ornamental flowers in China and have been cultivated all over the world. Yet the origin of chrysanthemum cultivars has been highly debated. In this study we employ the nuclear low-copy LFY gene to study the evolutionary history of chrysanthemum cultivars. The structure of the LFY gene in all Chrysanthemum species examined is highly conserved with three exons and two introns. The length of the LFY gene in Chrysanthemum varied from 2, 887 to 3, 348 bp. The two introns exhibited high levels of variation in length and sequence composition at the intraspecific and interspecific levels. Phylogenetic analysis of the whole LFY sequences of Chrysanthemum resulted in topologies that contained three major clades. The LFY sequences from the same cultivars are present in two or three clades, supporting that hybridization and allopolyploidy were important mechanisms in the origins of different chrysanthemums. Our results suggest that different cultivars had different ancestors. Chrysanthemum indicum, C. zawadskii and C. nankingense were likely the direct ancestors of most chrysanthemum cultivars examined. Chrysanthemum vestitum is a putative ancestor for some cultivars, and may have indirectly involved in the development of the chrysanthemum cultivars. Sequences of the LFY gene are informative to shed insights into the origin of chrysanthemum cultivars and show great potential as a phylogenetic marker to decipher the phylogeny of Chrysanthemum and its close relatives.

Analyzing the ploidy levels of plants is important for identifying species, selecting parental lines, identifying the relationships between species, and determining evolutionary patterns. The genus Chrysanthemum is widely distributed throughout the world and exhibits different ploidy levels. We used flow cytometry to analyze the ploidy levels of nine species of Chrysanthemum L. collected from different regions and geographical locations in China. Three diploids from Henan and Wuhan provinces corresponded to Chrysanthemum lavandulifolium and two species of C. nankingense, while three tetraploids from various regions corresponded to C. indicum and two species of C. chanetii. Two hexaploids corresponding to C. vestitum were collected at Funiu mountain (Henan province), and C. zawadskii was collected at Huangshan mountain (Anhui province). We found that OTTO extraction buffer was suitable for extracting nuclei from most species, apart from C. zawadskii. Flow cytometry proved to be a simple, rapid, and highly accurate method for identifying ploidy levels in Chrysanthemum species.

The multiobjective evolutionary algorithm (MOEA) based on decomposition transforms a multiobjective optimization problem into a set of aggregated subproblems and then optimizes them collaboratively. Since these subproblems usually have different degrees of difficulty, resource allocation (RA) strategies have been reported to enhance performance, attempting to dynamically assign proper amounts of computational resources for the solution of each of these subproblems. However, existing schemes for decomposition-based MOEAs fully rely on the relative improvement of the aggregated functions to do this. This paper proposes a diversity-enhanced RA strategy for this kind of MOEA, depending on both relative improvement on aggregated function value and solution density around each subproblem to assign computational resources. Thus, one subproblem surrounded with fewer solutions in its neighboring area and more relative improvement on the aggregated function value will be allocated a higher probability for evolution. Our experimental results show the advantages of our proposed strategy over two popular RA strategies available for decomposition-based MOEAs, on tackling a set of complicated benchmark problems.

The LEAFY (LFY) gene encodes a key transcription factor involved in flower development. In this study, we identified the LFY homolog in Chrysanthemum argyrophyllum L. (CaFL) and sequenced eight clones from a single plant. The cloned genes varied in length (2915–3075 bp) and each had three exons and two introns. The three exons were highly conserved, so intron variance was primarily responsible for gene length differences. The putative CaFL proteins from the eight clones were 99% identical to one another and greater than 97% identical to Chrysanthemum lavandulifolium DFL and Chrysanthemum nankingense CnFL. Intraspecific and interspecific sequence comparisons of the two introns indicated that they varied considerably in both length and sequence composition. Phylogenetic analysis showed that CaFL is more similar to dicotyledonous FLO/LFY-like proteins than to those from monocotyledons or gymnosperms.

Chrysanthemum is one of the most important commercially used floral crops. The flowering feature of chrysanthemum has considerable ecological and cultural value; however, it’s flowering mechanisms has not been well characterized to date because of a lack of genomic information. In this study, we firstly present extensive sequence and transcript abundance data for the transcriptome of a key chrysanthemum-related species, Chrysanthemum lavandulifolium (Fisch. ex Trautv), which was obtained using Illumina HiSeq™ 2000 sequencing platform. There were 108,737 high quality unigenes that were assembled based on 4 GB sequencing data, of which 58,093 were annotated by a similarity search with known proteins (E-value

AP1/FUL-like genes encode class-specific MADS box transcription factors in angiosperms and play important roles in the plant floral transition and floral organ identity. Chrysanthemum lavandulifolium (Fisch. ex Trautv.) Makino is one of the original species of Chrysanthemum. Understanding the function of AP1/FUL-like genes in C. lavandulifolium provide molecular-level insight into the flowering mechanism of Chrysanthemum. Here, an AP1/FUL-like gene was isolated from C. lavandulifolium. Sequence and phylogenetic analyses indicated that this gene, ClM8, belongs to the euFUL clade of the AP1/FUL-like genes. We observed different expression levels of ClM8 in C. lavandulifolium roots, stems, fruits and leaves and high expression in the flowers. A subcellular localisation assay showed that ClM8 is localised to the nucleus in vivo. The over-expression of ClM8 in Arabidopsis resulted in altered flowering time and inflorescence structure. These aberrant, transgenic Arabidopsis plants showed early flowering following the production of four to six small rosette leaves, lateral flowers and a composite terminal flower. The terminal flower consisted of two or four pistils surrounded by an abnormal number of sepals, petals and stamens. In addition, the morphology of the style of 35S::ClM8 plants silique differed from wild type, appearing thinner and longer. Further investigation suggested that the expression levels of FLOWERING LOCUS T (FT), LEAFY (LFY) and APETALA1 (AP1) were apparently increased in ClM8-transgenic Arabidopsis. Thus, we speculate that ClM8 promoted flowering in Arabidopsis due to the upregulation of FT. Taken together, our results suggest that ClM8 is involved in flowering induction and floral development that might also play a role in the development of fruits and leaves.

Leucanthemella linearis is an important marsh plant. The complete chloroplast genome sequence of L. linearis was obtained using next generation sequencing. It was 15,1401 bp in length, including a ...