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Precise nitrogen supply is crucial for ensuring the quality of cut chrysanthemums (Chrysanthemum morifolium Ramat.). The nitrogen nutrition index (NNI) serves as an important indicator for diagnosing crop nitrogen (N) nutrition. Hyperspectral remote sensing (HRS) technology has been widely used in monitoring crop N status due to its rapid, accurate, and non-destructive capabilities. However, its application in estimating the NNI of cut chrysanthemums has received limited attention. Therefore, this study aimed to use HRS to accurately determine the cut chrysanthemum NNI, thereby providing valuable guidance for managing N fertilization. During several key growth stages, a hyperspectral spectroradiometer was used to capture hyperspectral reflectance data (350–2500 nm) from three leaf layers. Subsequently, cut chrysanthemum canopies were sampled for aboveground biomass (AGB) and plant nitrogen concentration (PNC). The collected AGB and PNC data were then utilized to fit the critical N (Nc) dilution curve of cut chrysanthemums using a Bayesian hierarchical model, enabling the calculation of the NNI. Finally, spectral indices and partial least squares regression (PLSR) were used to establish the NNI estimation model for cut chrysanthemums. The results showed that the Nc dilution curve of the cut chrysanthemums was Nc = 5.401 × AGB−0.468. The first leaf layer (L1) proved to be optimal for estimating cut chrysanthemum NNI. Additionally, a newly proposed two-band spectral index, DVI-L1 (R1105, R700), demonstrated moderate predictive capabilities for the NNI of cut chrysanthemums (R2 = 0.5309, RMSE = 0.3210). Compared with the spectral index-based NNI estimation model, PLSR-L1 showed the best performance in estimating the cut chrysanthemum NNI (R2 = 0.8177, RMSE = 0.2000). Our results highlight the rapid NNI prediction potential of HRS and its significance in facilitating precise N management in cut chrysanthemums.

Crown diameter is one of the crucial indicators for evaluating the adaptability, growth quality, and ornamental value of garden chrysanthemums. To accurately obtain crown diameter, this study employed an unmanned aerial vehicle (UAV) equipped with a RGB camera to capture orthorectified canopy images of 64 varieties of garden chrysanthemums at different growth stages. Three methods, namely RGB color space, hue-saturation-value (HSV) color space, and the mask region-based convolutional neural network (Mask R-CNN), were employed to estimate the crown diameter of garden chrysanthemums. The results revealed that the Mask R-CNN exhibited the best performance in crown diameter estimation (sample number = 2409, R2 = 0.9629, RMSE = 2.2949 cm). Following closely, the HSV color space-based model exhibited strong performance (sample number = 2409, R2 = 0.9465, RMSE = 3.4073 cm). Both of the first two methods were efficient in estimating crown diameter throughout the entire growth stage. In contrast, the RGB color space-based model exhibited slightly lower performance (sample number = 1065, R2 = 0.9011, RMSE = 3.3418 cm) and was only applicable during periods when the entire plant was predominantly green. These findings provide theoretical and technical support for utilizing UAV-based imagery to estimate the crown diameter of garden chrysanthemums.

Nitrogen (N), phosphorus (P), and potassium (K) are three macronutrients that are crucial in plant growth and development. Deficiency or excess of any or all directly decreases crop yield and quality. There is increasing awareness of the importance of rhizosphere microorganisms in plant growth, nutrient transportation, and nutrient uptake. Little is known about the influence of N, P, and K as nutrients for the optimal production of Chrysanthemum morifolium. In this study, a field experiment was performed to investigate the effects of N, P, and K on the growth, nutrient use efficiency, microbial diversity, and composition of C. morifolium. Significant relationships were evident between N application rates, C. morifolium nutrient use, and plant growth. The N distribution in plant locations decreased in the order of leaf > stem > root; the distributions were closely related to rates of N application. Total P fluctuated slightly during growth. No significant differences were found between total P in the roots, stems, and leaves of C. morifolium vegetative organs. Principle component analysis revealed that combinations of N, P, and K influenced soil nutrient properties through their indirect impact on operational taxonomic units, Shannon index, and abundance of predominant bacterial taxa. Treatment with N, P, and K (600, 120, and 80 mg·plant−1, respectively) significantly improved plant growth and quality and contributed to the bacterial richness and diversity more than other concentrations of N, P, and K. At the flowering time, the plant height, leaf fresh weight, root dry weight, stem and leaf dry weight were increased 10.6%, 19.0%, 40.4%, 27% and 34.0%, respectively, when compared to the CK. The optimal concentrations of N, P, and K had a positive indirect influence on the available soil nutrient content and efficiency of nutrient use by plants by increasing the abundance of Proteobacteria, decreasing the abundance of Actinobacteria, and enhancing the potential functions of nitrogen metabolism pathways. N, P, and K fertilization concentrations of 600, 120, and 80 mg·plant−1 were optimal for C. morifolium cultivation, which could change environmental niches and drive the evolution of the soil microbial community and diversity. Shifts in the composition of soil microbes and functional metabolism pathways, such as ABC transporters, nitrogen metabolism, porphyrin, and the metabolism of chlorophyll II, glyoxylate, and dicarboxylate, greatly affected soil nutrient cycling, with potential feedback on C. morifolium nutrient use efficiency and growth. These results provide new insights into the efficient cultivation and management of C. morifolium.

The transcription factor family RELATED to ABSCISIC ACID INSENSITIVE3 (ABI3)/VIVIPAROUS1(VP1) (RAV) is a plant-specific group of transcription factors that only contain a conserved B3 DNA binding domain or both their own B3 and APETALA2 (AP2) domains belonging to the B3 superfamily, which is vital for plant growth, development, and stress response. Although genome-wide characterization and analysis of the RAV family genes have been conducted in some species, they have not been systematically reported in chrysanthemums. Here, we found six RAV family genes in the diploid Chrysanthemum seticuspe genome. Based on domain similarity and homology comparison analyses, RAV genes in Chrysanthemum were categorized into two clades: Class-I and Class-II. Conserved motif analysis revealed that all CsRAV proteins contained the B3 repression domain. An analysis of cis-acting elements suggested that CsRAV family genes may play parts in light, hormonal, abiotic stress, growth, and developmental processes. Furthermore, quantitative RT-PCR analysis validated that all six CsRAV genes responded to ethylene treatment, whereas the genes in the Class-I clade responded most significantly to ethylene. In summary, the above results provided a conceptual basis for further investigation into the functions of CsRAV genes in C. seticuspe.

The chrysanthemum is widely used as a cut flower, potted flower, and garden flower worldwide and has high ornamental, edible, and medicinal value. The flower heads, composed of ray florets and disc florets, are the most diverse in terms of morphology among ornamental plants. Here, we compared and analyzed the developmental processes of different capitulum types as well as ray florets and disc florets. Morphological differentiation of the two florets occurred on the dorsal domain of the petals at stage Ⅳ of flower development, and differences in stamen development occurred at stage Ⅴ. The dorsal domain of the ray florets and the early stage of flower development were also an essential site and period, respectively, for the differences among capitulum types. In situ hybridization revealed that CmCYC2c, whose homologs are involved in the specification of floret identity in Asteraceae, was expressed in both the dorsal and ventral domains of the ray petals in the tubular-type chrysanthemum, whereas, it was differentially transcribed in the ray petals of flat- and spoon-type chrysanthemum cultivars and had lower or no expression in the dorsal domain and higher expression in the ventral domain at stage Ⅳ. Our study indicates that the expression pattern of CmCYC2c on the dorsal domain of the ray floret at stage Ⅳ contributes to the formation of diverse flower head types in chrysanthemums.

The endophytic microbiomes significantly differed across tea chrysanthemum cultivars and organs (stems and leaves). The most abundant endophytic bacterial genera were Pseudomonas, Masillia, and Enterobacter in the leaves and Sphingomonas and Curtobacterium in the stems of the five cultivars. Meanwhile, the most abundant endophytic fungal genera in the leaves and stems of the five tea chrysanthemums were Alternaria, Cladosporium, and Sporobolomyces. Specifically, Rhodotorula was dominant in the leaves of ‘Jinsi huangjv’ and Paraphoma was dominant in the stems of ‘Jinsi huangjv’. In all cultivars, the diversity and richness of endophytic bacteria were higher in leaves than in stems (p < 0.05). The highest diversity and richness of endophytic bacteria were recorded in ‘Chujv’, followed by ‘Jinsi huangjv’, ‘Fubai jv’, ‘Nannong jinjv’, and ‘Hangbai jv’. Meanwhile, endophytic fungi were less pronounced. Twenty-seven and 15 cultivable endophytic bacteria and fungi were isolated, four isolated endophytic bacteria, namely, CJY1 (Bacillus oryzaecorticis), CY2 (Pseudomonas psychrotolerans), JSJ7, and JSJ17 (Enterobacter cloacae) showed higher indole acetic acid production ability. Further field studies indicated that inoculation of these four endophytic bacteria not only promoted plant growth and yield but also increased total flavonoids, chlorogenic acid, luteolin, and 3,5-dicoffeylquinic acid levels in the dry flowers of tea chrysanthemums.

Ethylene is a gaseous phytohormone that delays flowering in Chrysanthemum morifolium Ramat. To date, however, there have been no systematic studies on genes involved in the ethylene response of this species, and the mechanism underlying ethylene-delayed flowering remains unclear. Herein, we applied RNA sequencing to characterize the ethylene response by comparing the transcriptomes of chrysanthemum cultivar ‘Jinba’ with or without ethephon treatment. Six unique RNA-seq libraries were generated. The identified differentially expressed genes (DEGs) primarily involved ethylene, auxin, and abscisic acid signaling genes; circadian clock genes; genes encoding functional proteins associated with floral transition, such as homologs of AP1/FRUITFUL-like 1 (AFL1), TERMINAL FLOWER 1 (TFL1), and so on; and genes encoding transcription factors, specifically of the MYB and bHLH families. Furthermore, quantitative RT-PCR was used to verify the DEGs identified by RNA-seq. Heterologous CmAFL1 overexpression in Arabidopsis thaliana resulted in early flowering. Our findings present a landscape of transcriptomes and reveal the candidate genes involved in the ethylene-mediated regulation of flowering time in chrysanthemum, providing useful data for further studies.

As a molecular chaperone, HSP70 is widely involved in complex activities in plants. Under high temperature, drought, high salt, low temperature, heavy metals, and other stresses, HSP70 is rapidly synthesizes, stabilizes protein and biological macromolecular structures, and improves the stress resistance of plants. In this study, 83 ClHSP70 genes in Chrysanthemum lavandulifolium were identified based on the published Chrysanthemum lavandulifolium genome database. The genes were divided into six clusters based on a phylogenetic analysis, and the gene structures, conserved motifs, and functional domains were relatively conserved. Only two collinear genes were observed, and they formed a pair of duplicating genes. Multiple abiotic stress and phytohormone response elements were observed on the ClHSP70 promoter, such as temperature, drought, methyl jasmonate, abscisic acid, and other stress-related elements, and CpG islands were found on some ClHSP70 promoters, suggesting that they may be related to methylation modifications. Tissue expression analysis showed that the ClHSP70 genes were highly expressed in the roots. In addition, the gene expression changed significantly after 1 h of heat treatment and plays an important role in plant responses to temperature stress. The results of this study provide insights on the ClHSP70 gene family members and a theoretical basis for further research on functional analyses of the ClHSP70 gene family.

Nuclear factor Y (NF-Y) is a class of transcription factors (TFs) with various biological functions that exist in almost all eukaryotes. In plants, heterotrimers composed of different NF-Y subunits are numerous and have different functions that can participate in the regulation of plant growth at various stages. However, NF-Y genes have not been systematically analyzed in chrysanthemum, only involving several NF-Y members. In this study, forty-six NF-Y members were identified from the diploid species Chrysanthemum seticuspe, including eight NF-YA members, twenty-one NF-YB members, and seventeen NF-YC members. These NF-Y genes were analyzed for their physicochemical characteristics, multiple alignments, conserved motifs, gene structure, promoter elements, and chromosomal location. Phylogenetic analysis revealed that only two gene pairs in C. seticuspe underwent gene duplication events. The Ka/Ks ratios were both less than one, indicating that the two pairs underwent purifying selection. Promoter element analysis showed that multiple abiotic stress and hormone response elements were present in the CsNF-Y genes, suggesting that these genes play an important role in the response to stress, growth, and development in plants. Further validation of candidate genes in response to drought regulation using RT-qPCR demonstrated that CsNF-Y genes in C. seticuspe play an important role in drought regulation.

Chrysanthemummorifolium is one of the four major cut flowers in the world, with high ornamental and economic value. Flowering time is an important ornamental characteristic of chrysanthemum that affects its value in the market. In Arabidopsis, the FRUITFULL (FUL) gene plays a key role in inducing flowering. Here, we isolated an FUL clade MADS-box gene, CmFL3, from chrysanthemum inflorescence buds. CmFL3 localized in the cellular membrane and nucleus, and showed no transcriptional activity in yeast. The qRT-PCR assay showed that CmFL3 was strongly expressed in the leaves, receptacles, and disc floret petals. Furthermore, CmFL3 was mainly detected in the inflorescence meristem and bract primordia using in situ hybridization. Similar to Arabidopsis, overexpression of CmFL3 in chrysanthemum induced early flowering. Particularly, the expression level of CmAFT was downregulated, whereas that of CmFTL3 was upregulated in the leaves of transgenic chrysanthemum lines. Meanwhile, the overexpression of CmFL3 in Arabidopsis also led to earlier flowering. Furthermore, the expression of AtFT, AtAP1, AtLFY, and AtFUL was significantly increased in CmFL3 transgenic Arabidopsis. The present study verified the function of CmFL3 in regulating flowering time and further revealed that it could affect the expression of other flowering-related genes—CmAFT and CmFTL3. Therefore, the CmFL3 gene may be an important candidate for genetic breeding aimed at regulating flowering.

Transcription factors, also known as trans-acting factors, balance development and stress responses in plants. Branching plays an important role in plant morphogenesis and is closely related to plant biomass and crop yield. The apical meristem produced during plant embryonic development repeatedly produces the body of the plant, and the final aerial structure is regulated by the branching mode generated by axillary meristem (AM) activities. These branching patterns are regulated by two processes: AM formation and axillary bud growth. In recent years, transcription factors involved in regulating these processes have been identified. In addition, these transcription factors play an important role in various plant hormone pathways and photoresponses regulating plant branching. In this review, we start from the formation and growth of axillary meristems, including the regulation of hormones, light and other internal and external factors, and focus on the transcription factors involved in regulating plant branching and development to provide candidate genes for improving crop architecture through gene editing or directed breeding.

Garden chrysanthemums are a group of chrysanthemums (Chrysanthemum morifolium) that are mostly used in garden landscape collocations. Because most garden chrysanthemum varieties have low plants and poor space effects, they have certain limitations in garden landscape applications. In this study, we selected seven garden chrysanthemum varieties with excellent characteristics for artificial hybridization and intercross parentage to obtain new varieties with good traits we need. The phenotypic characteristics of the F1 offspring in terms of plant height, crown width, crown height ratio, number of main branches, branching intensity, plant form, inflorescence diameter, number of flowers per plant, number of ray florets, flower color, and days from planting period to coloring period were analyzed and systematically evaluated. Subsequently, a comprehensive evaluation system was established using the analytic hierarchy process (AHP) and k-means clustering method. We built a comprehensive analysis model, calculated the weighting of each evaluation factor, and multiplied the weight value by the score of the evaluation index standard of each factor to obtain the comprehensive score of each plant. All F1 plants were divided into four grades: excellent grade accounting for 13%, good grade accounting for 28%, medium grade accounting for 36%, and poor grade accounting for 23%. Then, we analyzed the differences of some quantitative traits between Group E (hybrids in excellent grade) and Group O (hybrids in good, medium, and poor grades). There were significant differences in plant height, crown height ratio, and the number of main branches but no significant difference in crown width. Combining with comprehensive score showed that Group E performed well overall. Finally, we selected five hybrid offspring with the highest overall scores in Group E as excellent variety materials of garden chrysanthemum for hedgerow. They were CH22, YQ73, HY08, CQ80, and HY07, respectively. We also found that plant height, lodging resistance, crown height ratio, plant form, crown width, and the number of main branches could be the main indicators in the AHP, which can be effectively applied to the comprehensive evaluation and breeding of garden chrysanthemums for hedgerows.

Heat shock proteins (HSPs) play important roles in various stress conditions. In this study, CmHSP90.5, whose expression is induced by heat and salt, was cloned from a chrysanthemum (Chrysanthemum morifolium) ‘Jinba’ and expressed in Arabidopsis. We found that CmHSP90.5 localized in the chloroplast. The heterologous expression of CmHSP90.5 weakened the heat tolerance of Arabidopsis and reduced the activity of antioxidant enzymes, including superoxide dismutase (SOD), catalase (CAT), and ascorbate peroxidase (APX), indicating that CmHSP90.5-mediated heat stress sensitivity may be partially due to the regulation of active oxygen cleavage. The levels of expression of AtHSP101, AtHSP15.7, and AtHSP17.6C in CmHSP90.5-overexpressing plants decreased compared with those in wild-type (WT) plants under heat stress, indicating that these HSPs and CmHSP90.5 coregulate a plant’s heat stress tolerance. In addition, the salt stress tolerance of the CmHSP90.5-overexpressing Arabidopsis decreased compared with that of WT plants; CmHSP90.5-overexpressing plants showed increased Na+ levels and decreased K+ and proline levels compared with those of WT plants. Interestingly, the expression of stress-related genes, such as the Na+/H+ antiporter encoding gene SOS1, high-affinity K+ transporter encoding gene HKT1;1, and proline synthesis gene AtP5CS1, decreased in CmHSP90.5-overexpressing plants under salt stress compared with those expressions in WT plants. Our findings lay a foundation for understanding the roles of CmHSP90.5 in response to abiotic stresses in chrysanthemum.

The architecture of spray cut chrysanthemum is crucial for the quality and quantity of cut flower production. However, the mechanism underlying plant architecture still needs to be clarified. In this study, we measured nine architecture-related traits of 195 spray cut chrysanthemum varieties during a two-year period. The results showed that the number of upper primary branches, number of lateral flower buds and primary branch length widely varied. Additionally, plant height had a significant positive correlation with number of leaf nodes and total number of lateral buds. Number of upper primary branches had a significant negative correlation with primary branch diameter, primary branch angle and primary branch length. Plant height, total number of lateral buds, number of upper primary branches, stem diameter, primary branch diameter and primary branch length were vulnerable to environmental impacts. All varieties could be divided into five categories according to cluster analysis, and the typical plant architecture of the varieties was summarized. Finally, a genome-wide association study (GWAS) was performed to find potential functional genes.

Chrysanthemum is a popular ornamental plant with a long history of cultivation. Both the leaf and flowerhead of Chrysanthemum are known to produce diverse secondary metabolites, particularly terpenoids. Here we aimed to determine the relationship between terpene chemistry and the trichome traits in Chrysanthemum. In our examination of three cultivars of C. morifilium and three accessions of C. indicum, all plants contained T-shaped trichomes and biseriate peltate glandular trichomes. The biseriate peltate glandular trichome contained two basal cells, two stalk cells, six secondary cells and a subcuticular space, while the non-glandular T-shaped trichome was only composed of stalk cells and elongated cells. Histochemical staining analysis indicated that the biseriate peltate glandular trichome contained terpenoids and lipid oil droplets but not the T-shaped trichome. Next, experiments were performed to determine the relationship between the accumulation and emission of the volatile terpenoids and the density of trichomes on the leaves and flowerheads in all six Chrysanthemum cultivars\accessions. A significant correlation was identified between the monoterpenoid and sesquiterpenoid content and the density of glandular trichomes on the leaves, with the correlation coefficients being 0.88, 0.86 and 0.90, respectively. In contrast, there was no significant correlation between the volatile terpenoid content and the density of T-shaped trichomes on the leaves. In flowerheads, a significant correlation was identified between the emission rate of terpenoids and the number of glandular trichomes on the disc florets, with a correlation coefficient of 0.95. Interestingly, the correlation between the density of glandular trichomes and concentrations of terpenoids was insignificant. In summary, the relationship between trichomes and terpenoid chemistry in Chrysanthemum is clearly established. Such knowledge may be helpful for breeding aromatic Chrysanthemum cultivars by modulating the trichome trait.

Plant-specific TCP transcription factors play a key role in plant development and stress responses. Chrysanthemum nankingense shows higher cold tolerance than its ornamental polyploid counterpart. However, whether the TCP gene family plays a role in conferring cold tolerance upon C. nankingense remains unknown. Here, we identified 23 CnTCP genes in C. nankingense, systematically analyzed their phylogenetic relationships and synteny with TCPs from other species, and evaluated their expression profiles at low temperature. Phylogenetic analysis of the protein sequences suggested that CnTCP proteins fall into two classes and three clades, with a typical bHLH domain. However, differences between C. nankingense and Arabidopsis in predicted protein structure and binding sites suggested a unique function of CnTCPs in C. nankingense. Furthermore, expression profiles showed that expression of most CnTCPs were downregulated under cold conditions, suggesting their importance in plant responses to cold stress. Notably, expression of miR319 and of its predicted target genes, CnTCP2/4/14, led to fast responses to cold. Overexpression of Arabidopsis CnTCP4 led to hypersensitivity to cold, suggesting that CnTCP4 might play a negative role in C. nankingense responses to cold stress. Our results provide a foundation for future functional genomic studies on this gene family in chrysanthemum.

Whole genome duplication, associated with the induction of widespread genetic changes, has played an important role in the evolution of many plant taxa. All extant angiosperm species have undergone at least one polyploidization event, forming either an auto- or allopolyploid organism. Compared with allopolyploidization, however, few studies have examined autopolyploidization, and few studies have focused on the response of genetic changes to autopolyploidy. In the present study, newly synthesized C. nankingense autotetraploids (Asteraceae) were employed to characterize the genome shock following autopolyploidization. Available evidence suggested that the genetic changes primarily involved the loss of old fragments and the gain of novel fragments, and some novel sequences were potential long terminal repeat (LTR) retrotransposons. As Ty1-copia and Ty3-gypsy elements represent the two main superfamilies of LTR retrotransposons, the dynamics of Ty1-copia and Ty3-gypsy were evaluated using RT-PCR, transcriptome sequencing, and LTR retrotransposon-based molecular marker techniques. Additionally, fluorescence in situ hybridization(FISH)results suggest that autopolyploidization might also be accompanied by perturbations of LTR retrotransposons, and emergence retrotransposon insertions might show more rapid divergence, resulting in diploid-like behaviour, potentially accelerating the evolutionary process among progenies. Our results strongly suggest a need to expand the current evolutionary framework to include a genetic dimension when seeking to understand genomic shock following autopolyploidization in Asteraceae.

The production of chrysanthemums is severely hampered by Fusarium wilt, which is exacerbated by monoculture. In this study, the role of inorganic plant nutrition fertilizer (IPN), organic fertilizer (OF) and bio-organic fertilizer (BOF) in avoiding monoculture-related production constraints was evaluated. We conducted a series of greenhouse experiments and studied the growth of chrysanthemum and changes in rhizosphere soil microflora and function. BOF application reduced the incidence of Fusarium wilt by 82.8% and increased the chrysanthemum shoot height and flower ray floret number by 31.4% and 26.1%, respectively. High-throughput Illumina HiSeq2500 sequencing results indicated that BOF and OF treatments increased the values of α-diversity indices of bacteria and fungi. In addition, significant alterations in microbe community structures were found in response to IPN, OF or BOF application. Among the major genera detected after BOF treatments, the levels of Fusarium and Glycomyces decreased while Cladosporium, arbuscular mycorrhizal and endophyte groups increased. In particular, the abundance of Mariniflexile had a positive relationship (R = 0.693, p < 0.05) with the incidence of Fusarium wilt, while Cladosporium showed a significant negative relationship (R = −0.586, p < 0.05). Interestingly, an analysis of microbiomes based on 16S rRNA sequences revealed that the functions of signal transduction, bacterial secretion system, oxidative phosphorylation and the metabolism of carbohydrate, nitrogen and amino acids all increased in both BOF and OF treatments. The results suggested that BOF could be effective for chrysanthemum monoculture soil restoration, potentially by altering the microbial community structures and functions, which affect the physiological and morphological attributes of chrysanthemum in monoculture.

Chlorophyll is vital for photosynthesis to produce sugars and other useful biochemical products in green plants. However, the molecular effects of chlorophyll deficiency in Chrysanthemum are largely unknown. In this study, we identified a bud sport mutant chrysanthemum belonging to the variety ‘Nannong Binyun’, which has yellow branches. Plant physiological studies have shown that the yellow color is revealed due to chlorophyll loss. RNA extracts of yellow and green tissues were analyzed using high-throughput RNA-sequencing, and a total of 11,649 tissue enriched unigenes that respond to chlorophyll deficiency were identified, including 4803 unigenes upregulated in yellow tissues and 6846 unigenes in green tissues. GO analysis revealed that these tissue-enriched genes may involve in the physiological processes of chlorophyll accumulation and photosynthesis. In addition, many DEGs from the families of AP2-EREBP, bHLH, MYB, and FAR1 that are associated with plant development and stress response were detected. Our study found that most of the genes from the GRAS family were downregulated in yellow leaves, indicating their putative roles in stem cell maintenance and possible contribution to leaf size determination.

Decapitation is common in horticulture for altering plant architecture. The decapitation of chrysanthemum plants breaks apical dominance and leads to more flowers on lateral branches, resulting in landscape flowers with good coverage. We performed both third- and second-generation transcriptome sequencing of the second buds of chrysanthemum. This third-generation transcriptome is the first sequenced third-generation transcriptome of chrysanthemum, revealing alternative splicing events, lncRNAs, and transcription factors. Aside from the classic hormones, the expression of jasmonate-related genes changed because of this process. Sugars also played an important role in this process, with upregulated expression of sucrose transport-related and TPS genes. We constructed a model of the initial growth of the second buds after decapitation. Auxin export and sugar influx activated the growth of these buds, while the JA-Ile caused by wounding inhibited the expression of CycD genes from 0 h to 6 h. After wound recovery, cytokinins accumulated in the second buds and might have induced ARR12 expression to upregulate CycD gene expression from 6 h to 48 h, together with sugars. Therefore, jasmonates, cytokinins, sugars, and auxin work together to determine the fate of the buds of plants with short internodes, such as chrysanthemum.

Roots provide anchorage and enable the absorption of water and micronutrients from the soil for plants. Besides these essential functions, roots are increasingly being recognized as an important organ for the production of diverse secondary metabolites. The goal of this study was to investigate the chemical composition and function of terpenoid secondary metabolites in roots of different cultivars of the popular ornamental plant Chrysanthemum morifolium Ramat. Although C. morifolium is known for rich production of secondary metabolites in its flower heads and leaves, the diversity of secondary metabolites in roots remains poorly characterized. In this study, 12 cultivars of C. morifolium were selected for comparative analysis. From their roots, a total of 20 terpenoids were detected, including four monoterpenes, 15 sesquiterpenes, and one diterpene. The cultivar ‘She Yang Hong Xin Ju’ exhibited the highest concentration of total terpenoids at approximately 730 µg·g−1 fresh weight. Most cultivars contained sesquiterpenes as the predominant terpenoids. Of them, (E)-β-farnesene was detected in all cultivars. Based on their terpenoid composition, the 12 cultivars were planed into four groups. To gain insights into the function of root secondary metabolites, we performed bioassays to assess their effects on growth of three species of pathogenic fungi: Fusarium oxysporum, Magnaporthe oryzae, and Verticillium dahliae. Significant variability in antifungal activity of the root extracts among different cultivars were observed. The cultivar ‘Xiao Huang Ju’ was the only cultivar that had significant inhibitory effects on all three species of fungi. Our study reveals the diversity of terpenoids in roots of C. morifolium and their function as a chemical defense against fungi.

The objective of this study was to identify the major volatile compounds and their relative concentrations in flowers of different chrysanthemum cultivars and their wild relatives. The volatile organic components of fresh flowers were analyzed using a headspace solid-phase microextraction coupled with gas chromatography-mass spectrometry. In total, 193 volatile organic components were detected; the major scent components were monoterpenoids and oxygenated monoterpenoids, which accounted for 68.59%–99.93% of the total volatiles in all tested materials except for Chrysanthemum indicum collected from Huangshan, in which they accounted for only 37.45% of total volatiles. The major volatile compounds were camphor, α-pinene, chrysanthenone, safranal, myrcene, eucalyptol, 2,4,5,6,7,7ab-hexahydro-1H-indene, verbenone, β-phellandrene and camphene. In a hierarchical cluster analysis, 39 accessions of Chrysanthemum and its relatives formed six clusters based on their floral volatile compounds. In a principal component analysis, only spider type flowers were located closely on the score plot. The results of this study provide a basis for breeding chrysanthemum cultivars which desirable floral scents.

Chrysanthemum morifolium Ramat is an ornamental plant of worldwide cultivation. Like many other species in the family Asteraceae, C. morifolium is a rich producer of secondary metabolites. There are two objectives in this study: (I) to determine and compare the diversity of apolar secondary metabolites among different cultivars of C. morifolium and (II) to compare their properties as antifungal agents. To attain these objectives, we selected 13 cultivars of C. morifolium that are commonly used for making chrysanthemum tea as experimental materials. Leaves at the same developmental stage were collected from respective mature plants and subjected to organic extraction. The extracts were analyzed using gas chromatography−mass spectrometry. A total of 37 apolar secondary metabolites including 26 terpenoids were detected from the 13 cultivars. These 13 cultivars can be largely divided into three chemotypes based on chemical principal components analysis. Next, the extracts from the 13 cultivars were examined in in vitro assays for their antifungal properties against three species of pathogenic fungi: Fusarium oxysporum, Magnaporthe oryzae, and Verticillium dahliae. Significant variability in antifungal activity of the leaf extracts among different cultivars was observed. The 13 cultivars can be divided into four groups based on their antifungal activities, which could be partly correlated to the contents of terpenoids. In short, this study reveals large variations in chemical composition, particularly of terpenoids, of leaf secondary metabolites among different cultivars of C. morifolium and their different abilities in functioning as antifungal agents.

Chemical fungicides are a frequently used intervention for the control of the Fusarium wilt of chrysanthemum, but are no longer considered environmentally friendly. However, the biofungicides offer one of the best alternatives to reduce the use of chemical fungicides. In this study, a series of two-year greenhouse experiments were conducted to evaluate the soil-applied chemical fungicide (dazomet, DZ) and biofungicide (biocontrol agent combined with B. subtilis NCD-2, BF) for controlling the Fusarium wilt of chrysanthemum and its effects on rhizosphere soil microbiota. The results indicated that DZ and BF showed good control efficacy of Fusarium wilt of chrysanthemum in the two-year application evaluation. However, the DZ treatment significantly decreased the soil catalase and urease activities compared with the control, while BF showed a significant increase in bacterium/fungus ratios (B/F), soil urease and acid phosphatase activities. Abundances of potential plant pathogens F. oxysporum, Rhizoctonia zeae and Rhizoctonia solani were also lower, while potential plant-growth-promoting micro-organisms like the Rhizobiales bacterium and Mariniflexile sp. were higher in the BF treatment than in the control. Our findings suggested that the overall effect of the soil biota on chrysanthemum growth was more positive and stronger in the BF treatment than in the DZ treatment.

The green organs, especially the leaves, of many Compositae plants possess characteristic aromas. To exploit the utility value of these germplasm resources, the constituents, mainly volatile compounds, in the leaves of 14 scented plant materials were qualitatively and quantitatively compared via gas chromatography-mass spectrometry (GC-MS). A total of 213 constituents were detected and tentatively identified in the leaf extracts, and terpenoids (especially monoterpene and sesquiterpene derivatives), accounting for 40.45–90.38% of the total compounds, were the main components. The quantitative results revealed diverse concentrations and compositions of the chemical constituents between species. Principal component analysis (PCA) showed that different groups of these Compositae plants were characterized by main components of α-thujone, germacrene D, eucalyptol, β-caryophyllene, and camphor, for example. On the other hand, cluster memberships corresponding to the molecular phylogenetic framework, were found by hierarchical cluster analysis (HCA) based on the terpenoid composition of the tested species. These results provide a phytochemical foundation for the use of these scented Compositae plants, and for the further study of the chemotaxonomy and differential metabolism of Compositae species.

Sustained monoculture often leads to a decline in soil quality, in particular to the build-up of pathogen populations, a problem that is conventionally addressed by the use of either fungicide and/or soil fumigation. This practice is no longer considered to be either environmentally sustainable or safe. While the application of organic fertilizer is seen as a means of combating declining soil fertility, it has also been suggested as providing some control over certain soil-borne plant pathogens. Here, a greenhouse comparison was made of the Fusarium wilt control efficacy of various treatments given to a soil in which chrysanthemum had been produced continuously for many years. The treatments comprised the fungicide carbendazim (MBC), the soil fumigant dazomet (DAZ), the incorporation of a Paenibacillus polymyxa SQR21 (P. polymyxa SQR21, fungal antagonist) enhanced bio-organic fertilizer (BOF), and applications of BOF combined with either MBC or DAZ. Data suggest that all the treatments evaluated show good control over Fusarium wilt. The MBC and DAZ treatments were effective in suppressing the disease, but led to significant decrease in urease activity and no enhancement of catalase activity in the rhizosphere soils. BOF including treatments showed significant enhancement in soil enzyme activities and microbial communities compared to the MBC and DAZ, evidenced by differences in bacterial/fungi (B/F) ratios, Shannon–Wiener indexes and urease, catalase and sucrase activities in the rhizosphere soil of chrysanthemum. Of all the treatments evaluated, DAZ/BOF application not only greatly suppressed Fusarium wilt and enhanced soil enzyme activities and microbial communities but also promoted the quality of chrysanthemum obviously. Our findings suggest that combined BOF with DAZ could more effectively control Fusarium wilt disease of chrysanthemum.