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Lilies ( Lilium spp.) are renowned for their diverse and captivating floral scents, which are highly valued both commercially and ornamentally. This review provides a comprehensive overview of recent advancements in the identification, biosynthesis, and regulation of fragrance components in lily flowers. Various volatile organic compounds (VOCs) that contribute to the unique scents of different lily species and cultivars, including terpenoids, benzenoids/phenylpropanoids, and fatty acid derivatives, are discussed. The release patterns of these compounds from different floral tissues and at different developmental stages are examined, highlighting the significant role of tepals. Detection methods such as gas chromatography-mass spectrometry (GC-MS) and sensory analysis are evaluated for their effectiveness in fragrance research. Additionally, the biosynthetic pathways of key fragrance compounds are explored, focusing on the terpenoid and benzenoid/phenylpropanoid pathways and the regulatory mechanisms involving transcription factors and environmental factors. This review also addresses the influence of genetic and environmental factors on fragrance production and proposes future research directions to enhance the aromatic qualities of lilies through selective genetic and breeding approaches. Emphasis is placed on the potential applications of these findings in the floral industry to improve the commercial value and consumer appeal of lily flowers.

Trichomes play a crucial role in plant resistance to abiotic and biotic stresses, and their development and characteristics vary across different species. This study demonstrates that trichomes of Lilium pumilum exhibit synchronized growth during flower bud differentiation and enhance the plant's adaptability to UV-B radiation and aphid infection. We identified LpNAC48, a NAC family transcription factor (TF), that interacted with the B-box (BBX) family TF LpBBX28, during trichome formation in L. pumilum. Silencing LpNAC48 or LpBBX28 impaired trichome development and reduced trichome density on the outer perianths. We demonstrated that the upstream regulators LpNAC48 and LpBBX28 directly bound to the promoter of the bHLH TF-encoding gene LpGL3-LIKE (LpGL3L) to activate its expression. Moreover, an ABA-responsive element within a 259-bp DNA variation in the LpNAC48 promoter was important for its expression and was bound by the bZIP TF LpbZIP29 during trichome development. This binding activated LpNAC48 expression and contributed to trichome formation. This study provides insights into the role of a small DNA sequence variation in gene expression and trichome traits., (© The Author(s) 2025. Published by Oxford University Press on behalf of American Society of Plant Biologists. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Gray mold caused by Botrytis elliptica is one of the most determinative factors of lily growth and has become a major threat to lily productivity. However, the nature of the lily B. elliptica interaction remains largely unknown. Here, comparative transcriptomic and metabolomic were used to investigate the defense responses of resistant ('Sorbonne') and susceptible ('Tresor') lily cultivars to B. elliptica infection at 24 hpi. In total, 1326 metabolites were identified in 'Sorbonne' and 'Tresor' after infection, including a large number of phenylpropanoids. Specifically, the accumulation of four phenylpropanes, including eriodictyol, hesperetin, ferulic acid, and sinapyl alcohol, was significantly upregulated in the B. elliptica -infected 'Sorbonne' compared with the infected 'Tresor', and these phenylpropanes could significantly inhibit B. elliptica growth. At the transcript level, higher expression levels of F3'M , COMT , and CAD led to a higher content of resistance-related phenylpropanes (eriodictyol, ferulic acid, and sinapyl alcohol) in 'Sorbonne' following B. elliptica infection. It can be assumed that these phenylpropanes cause the resistance difference between 'Sorbonne' and 'Tresor', and could be the potential marker metabolites for gray mold resistance in the lily. Further transcriptional regulatory network analysis suggested that members of the AP2/ERF, WRKY, Trihelix, and MADS-M-type families positively regulated the biosynthesis of resistance-related phenylpropanes. Additionally, the expression patterns of genes involved in phenylpropanoid biosynthesis were confirmed using qRT-PCR. Therefore, we speculate that the degree of gray mold resistance in the lily is closely related to the contents of phenylpropanes and the transcript levels of the genes in the phenylpropanoid biosynthesis pathway. Our results not only improve our understanding of the lily's resistance mechanisms against B. elliptica , but also facilitate the genetic improvement of lily cultivars with gray mold resistance.

Lily (Lilium asiatica) is an important plant grown for its range of flower colors and heavy scent. In March 2024, potyvirus-like symptoms consisting of light-yellow mottling and mosaic were noticed on 12/20 lily plants in a private property in Weslaco, Hidalgo County, Texas. Two symptomatic plants (WTX1 and WTX2) were sampled randomly for virus diagnosis. The leaf extracts of both samples were negative for the potyvirus group using Agdia's Poty ImmunoStrip® (Agdia, Inc., Elkhart, IN, USA). However, rub-inoculation of the extracts onto healthy Nicotiana benthamiana and Vigna unguiculata plants (n=4, each) induced mild mottle symptoms on the systemic leaves of both herbaceous test plants 20 to 28 days post inoculation, indicating the presence of a mechanically transmissible agent in the samples. No virus-like symptoms were observed on the mock-inoculated plants (n=1, each) of both species. To test for suspected potyvirus infection, 2-µg of total nucleic acid extracts (Dellaporta et al. 1983) from WTX1 and WTX2 were used for complimentary DNA (cDNA) synthesis with Oligo(dT) primer and the PrimeScript 1st strand cDNA synthesis kit (Takara Bio, USA). One microliter aliquot of each cDNA served as template in 12.5-µl conventional PCR reaction volumes with 5 U Taq polymerase (Roche Diagnostics, Indianapolis, IN), and two pairs of degenerate primers targeting the partial cylindrical inclusion (CI) gene and the helper component protease (HC-Pro) of potyviruses (Ha et al. 2008). The expected ~700-bp DNA product of each gene target was amplified from both samples. The amplicons were excised, gel eluted (Zymoclean™ Gel DNA Recovery kit) and cloned individually into the pJET1.2/Blunt vector (Life Technologies). Recombinant plasmids from two transformed Escherichia coli cells per cloned DNA insert were Sanger sequenced. BLASTn analysis of each gene-specific nucleotide (nt) sequence fragment revealed their identities (83 to 92 % nt) as lily mottle virus (LMoV; genus Potyvirus) at 93 to 98% query coverage. In pairwise comparison, the obtained sequences of LMoV isolates from TX specific to the CI (GenBank accession nos. PQ037260-61) and the HC-Pro (PQ037262-63) shared ~99% nt and 100% amino acid (aa) identities with each other and ~91-92% nt and 98-100% aa identities with the closest isolate, Bate5 (JN127341) from Australia. Based on these results, a pair of LMoV-specific primers (LM_v109-F: 5'-GGCCAGTAATGTGCACAAGC-3' and LM_c527-R: 5'-TCGCTGTAGCTAGCGACGTAC-3') was newly designed to target the partial CI gene, internal to the 700-bp fragment obtained above with the generic primers. The primer pair was used to screen each of the mechanically inoculated test plants by RT-PCR as previously described above. The expected 438-bp fragment of the LMoV CI gene was amplified from all the inoculated plants of both N. benthamiana and V. unguiculata; the results were confirmed by Sanger sequencing as described above. The mock-inoculated plant of each experimental host plant tested negative for LMoV. LMoV is a quarantine pest and was previously reported in the U.S. (Brierley and Smith, 1944) and other lily-producing parts of the world, such as the Netherlands, China, Korea, and Brazil. To our knowledge, this is the first confirmed report of the virus in Texas, thus expanding its geographical range. Testing of lily foundation stocks before propagation is essential to prevent further spread of LMoV via planting material.

Cotton aphids ( Aphis gossypii Glover) cause harm by feeding on phloem sap and spreading plant viruses to lily. Understanding the mechanisms by which aphids infest lily plants is crucial for effective aphid management and control. In this study, we investigated the activity of antioxidants, integrated nontargeted metabolomes and transcriptomes of lilies infested by cotton aphids to explore the changes in lily leaves. Overall, the results indicated that the catalase (CAT) activity in the leaves of the lily plants was greater than that in the leaves of the control plants. A comprehensive identification of 604 substances was conducted in the leaves. Furthermore, the differentially abundant metabolite analysis revealed the enrichment of phenylalanine metabolism and α-linolenic acid metabolism. Moreover, 3574 differentially expressed genes (DEGs), whose expression tended to increase, were linked to glutathione metabolism and phenylpropanoid biosynthesis. In addition, the integrated analysis revealed that the defensive response of lily leaves to aphids is manifested through antioxidant reactions, phenylpropane and flavonoid biosynthesis, and α-linolenic acid metabolism. Finally, the key metabolites were CAT, glutathione, coumaric acid, and jasmonic acid, along with the key genes chalcone synthase ( CHS ), phenylalanine ammonia-lyase ( PAL ), and 12-oxo-phytodienoic acid reductase ( OPR ). Accordingly, the findings of this research elucidate the molecular and metabolic reactions of A. gossypii in lily plants, offering valuable insights for developing aphid resistance strategies in lily farming.

As a typical bulb flower, lily is widely cultivated worldwide because of its high ornamental, medicinal and edible value. Although breeding efforts evolved over the last 10000 years, there are still many problems in the face of increasing consumer demand. The approach of biotechnological methods would help to solve this problem and incorporate traits impossible by conventional breeding. Target traits are dormancy, development, color, floral fragrance and resistances against various biotic and abiotic stresses, so as to improve the quality of bulbs and cut flowers in planting, cultivation, postharvest, plant protection and marketing. Genetic transformation technology is an important method for varietal improvement and has become the foundation and core of plant functional genomics research, greatly assisting various plant improvement programs. However, achieving stable and efficient genetic transformation of lily has been difficult worldwide. Many gene function verification studies depend on the use of model plants, which greatly limits the pace of directed breeding and germplasm improvement in lily. Although significant progress has been made in the development and optimization of genetic transformation systems, shortcomings remain. Agrobacterium-mediated genetic transformation has been widely used in lily. However, severe genotypic dependence is the main bottleneck limiting the genetic transformation of lily. This review will summarizes the research progress in the genetic transformation of lily over the past 30 years to generate the material including a section how genome engineering using stable genetic transformation system, and give an overview about recent and future applications of lily transformation. The information provided in this paper includes ideas for optimizing and improving the efficiency of existing genetic transformation methods and for innovation, provides technical support for mining and identifying regulatory genes for key traits, and lays a foundation for genetic improvement and innovative germplasm development in lily., (© 2024. The Author(s).)

The present study aimed to investigate the variations in the nutritional composition, antioxidant capacity, and metabolite profile of lilies subjected to different drying treatments, including vacuum freeze drying (VFD), hot air drying (HAD), vacuum drying (VD), and infrared drying (ID). The results show that VFD provided better preservation of the original coloration and displayed the highest levels of total amino acid content, total phenolic content, total flavonoid content, and polysaccharide and alkaloid content. Our results reveal that VFD treatment can be employed to obtain high-quality lilies with desirable appearance characteristics and nutrient compositions. Metabolomics analysis identified a total of 464 metabolites from various dried lilies. Differential metabolite screening found 150 differential metabolites across all pairwise comparisons. Hierarchical clustering analysis (HCA) indicated that lilies subjected to VFD treatment exhibited a higher abundance of steroids, saponin, flavonoids, and phenolic glycoside, whereas those subjected to HAD, VD, or ID treatments showed relatively elevated levels of specific amino acids or derivatives. This study elucidates the significant impact of various drying treatments on the quality and metabolic profile of lilies, thereby providing valuable insights for enhancing the nutritional quality of processed lilies.

The Lanzhou lily is a regionally distinctive vegetable; the emergence of lightly processed lilies has addressed the inconvenience of consuming fresh lilies. However, the cleaning and impurity removal during the processing of lightly processed lily may strip off its original protective barrier and affect the edible quality. As one of the preservation methods, tea tree essential oil (TTEO) has the characteristics of being green, safe, and efficient preservative properties. This study focused on investigating the effects of different concentrations (25 μL/L, 50 μL/L, and 100 μL/L) of TTEO on the quality and microbiological safety of lightly processed lily. The results showed that compared with the control, appropriate concentrations of TTEO treatment could delay weight loss, improve appearance, firmness, and sensory quality, and maintain microbiological safety with the best effect observed at 50 μL/L. Meanwhile, TTEO treatment induced phenylalanine ammonia-lyase activity, thereby increasing the total phenolic content. Furthermore, TTEO enhanced the superoxide dismutase (SOD) and ascorbate peroxidase (APX) activity, which reduced O 2 -· production rate and H 2 O 2 content. TTEO inhibited lipoxygenase (LOX) activity, reducing the relative conductivity and malondialdehyde content, thereby delaying lipid peroxidation and quality deterioration. This indicates that TTEO could enhance antioxidant capacity by regulating reactive oxygen species (ROS) metabolism and delay the quality deterioration of lightly processed lily by inhibiting lipid peroxidation.

To study the genetic background of lily ( Lilium spp.) germplasm resources, and accurately evaluate and select excellent germplasm for genetic improvement of lily, we analyzed the genetic background of 62 lily germplasm accessions from 11 provinces of China by using simple sequence repeat (SSR) molecular markers. The results showed that 15 out of 83 pairs of lily SSR primers were polymorphic. A total of 157 allelic loci were amplified, with the number of alleles per locus ranging from 5 to 19 and the average number of effective alleles per locus being 4.162 8. The average observed heterozygosity and expected heterozygosity were 0.228 2 and 0.694 1, respectively. The average polymorphic information content was 0.678 8. The average Nei's diversity index and Shannon's information index were 0.694 1 and 1.594 9, respectively, indicating that the tested lily germplasm had high genetic diversity. The 62 germplasm accessions were classified into 5 groups by the unweighted pair group method with arithmetic mean (UPGMA) and into 3 groups by the principal component analysis. The two analyses revealed a geographic correlation among different groups. The majority of lily germplasm accessions from the same source tended to cluster together. The population structure analysis classified the lily accessions into 4 populations and 1 mixed population. The above results provide a theoretical basis and genetic resources for the precise identification and breeding of lily germplasm resources.

Lily (Lilium spp.) is a valuable ornamental bulb flower plant in Liliaceae, and its bulbs have high edible and medicinal value. Compared with bulb propagation of other lilies, seed propagation and short growth period are the most significant characteristics of Lilium×formolongi. In 2023, leaf rot disease (LRD) was observed on approximately 70% of the Lilium×formolongi seedlings sown in an experimental greenhouse in Wuhan, Hubei province, China. Irregular brown water-soaked spots were discovered in the early stages of infected seedlings. Then, spots spread throughout the leaves and caused the leaves to brown, soften, and wilted. A pathogen associated with symptoms was isolated by incubating sterilized leaves on potato dextrose agar plates at 25 ℃ for 2-3 days. Then, a pure single colony was isolated through a single hyphal tip isolation method. The fungal colony was white with abundant aerial mycelium and produced a yellow pigment diffusible into the agar. Microscopically, isolated mycelia were reticulate and pale yellow, while conidia were dark brown, smooth, and spherical, 7.31 to 6.98 × 4.03 to 3.87μm (average 5.44×5.41μm; n=30); oval in lateral view, and had a light stripe in the middle. To identify the species of the fungus at the molecular level, ITS and EF-1α genes were amplified and sequenced using primers ITS1/ITS4 (M Gardes et al. 1993) and 758F/986R (Carbone and Kohn 1999). The BLAST results in GenBank showed that the ITS(OR523578) and EF-1α(PP066842) sequences of LRD shared 99.82% and 99.24% identity with the distinct Apiospora paraphaeosperma strains (GenBank accession MT040110, ON806628.1, respectively). Combined with the morphology of the colony and conidium, the fungus was identified as Ap. paraphaeosperma. In the pathogenicity test, six healthy leaves were inoculated with mycelium disc and then kept in an incubator (22 ℃, 90% humidity, 16h light /8h darkness). The inoculated leaves showed necrosis and wilt symptoms similar to those observed in the greenhouse, while the control leaves were asymptomatic. A re-isolation, morphology identification and DNA sequencing of the fungus confirmed its infection with Ap. paraphaeosperma in Lilium spp. At present, rot caused by Ap. paraphaeosperma has only been reported in Thailand and South Korea, both of which are found on bamboo stems (Hyde et al. 2016; Sun Lul Kwon et al. 2022). As far as we know, this is the first report of leaf rot of lily caused by Ap. paraphaeosperma in China. This report can help identify this disease and further develop effective control measures.

Lily ( Lilium brownii var. Viridulum Baker ) is a well-known edible plant with large, white and sweet bulb scales that has important medicinal value (Zhou et al. 2021) and is grown mainly in the Hebei, Shanxi and Henan provinces of China. In May 2021, a case of bulb rot was discovered in a 3.33 hm 2 plantation in Huaihua, Hunan Province, affecting 20% of the area (27°59'30″N, 110°32'20″E). The disease is most severe during the rainy season in May and June. In the early stage, irregular brown spots appeared on the lily scales, the necrosis was depressed and gradually enlarges, and in the later stage, the scales were scattered from the base of the disc and slough off. Ten samples were taken randomly from different plants in the plantation area to isolate the pathogens. After washing with sterile water, they were cut into small pieces and sterilised with 3% hydrogen peroxide for 30 s, 75% ethanol for 90 s, rinsed three times with sterile water and dried on sterile filter paper, then placed on a water agar plate and incubated in the dark in a constant temperature incubator at 28℃ for 3 to 5 days. After 2 days, the mycelium at the edge of the colony was transferred to a PDA plate and incubated for 3-5 days at 28°C in the dark to obtain pure fungal isolates. Eighteen purified fungal isolates were obtained, of which sixteen looked like Fusarium (88.9% isolation rate) and three representative isolates (BHBR2, BHBR3 and BHBR5) were selected for further study. The surface of this fungus was white with dense aerial mycelium. Some had an orange centre in the medium. Microconidia were oval in shape and appeared either straight or slightly curved. These microconidia were colourless, had 0-1 septa and measured 3.334 to 14.724 × 2.216 to 5.385 μm (n=100). Macroconidia were predominantly three-septate, crescent-shaped structures that were thin-walled and slightly curved. Cells at the apex and base were similarly curved. Macroconidia measured 17.956 to 32.150 × 2.788 to 4.492 μm (n=100). The mitochondrial small subunit (mtSSU) and translation elongation factor 1-α (TEF1) genes were amplified and sequenced using the NMS1/NMS2 and TEF-R/TEF-F primers to verify the identity of the pathogens (Stewart et al. 2006). The sequences were submitted to GenBank (BHBR2: mtSSU, PP273435; TEF, OR900976; BHBR3: mtSSU, PP277729; TEF, OR900977; BHBR5: mtSSU, PP277728; TEF, OR900978). A concatenated phylogenetic tree of the two genes was constructed and analysis showed that BHBR2, BHBR3 and BHBR5 were significantly clustered with Fusarium commune . Based on the results of morphological identification and phylogenetic tree analysis, the three isolates were identified as Fusarium commune . We carried out pathogenicity tests using two methods, one in which 6 × 6 mm fungal blocks were inoculated on lily ( L. brownie var. viridulum Baker ) scales and controls inoculated with sterile blocks, and the other in which strain BHBR2 was selected to carry out pathogenicity tests on bulbs of live plants soaked with 50 ml of a 1 × 10 6 conidial suspension and bulbs of control plants soaked with sterile water, all in three replicates. They were placed in a growth chamber at 28°C and 80% relative humidity, and the scales were moistened with moistened sterile filter paper. After 3 days of rearing treated scales, lesions appeared on lily scales inoculated with mycelial blocks and expanded with time, whereas no lesions appeared on lily scales inoculated with sterile blocks. One month later, whole plants soaked in the spore suspension wilted, while the control plants grew well. The pathogens re-isolated from the diseased tissues had the same morphological characteristics as representative isolates. This confirms Koch's hypothesis. Fusarium commune has been shown to be the most important pathogenic fungus causing root rot in Alfalfa ( Medicago sativa ) (Yang et al. 2022) and blueberry ( Vaccinium uliginosum L. ) (Li et al. 2023) in China. To our knowledge, this is the first report of Fusarium commune causing lily bulb rot in the world, which will lay the foundation for future control of lily bulb rot.

Objective: Lily is an essential ornamental flowering species worldwide. Drought stress is a major constraint affecting the morphology and physiology and lily leaves and flowers. Therefore, understanding the molecular mechanism underlying lily response to drought stress is important., Method: Transcriptome and metabolome analysis were performed on Oriental Lily subjected to drought stress., Result: Most transcription factors and metabolites yielded by the conjoint analysis displayed a downregulated expression pattern. Differential genes and metabolites mainly co-enriched in glycolic pathways related to sugars, such as galactose, and sucrose, glycolysis and gluconeogenesis, indicating that drought stress reduced the sugar metabolism level of Oriental Lily. Combined with transcriptome and metabolome data, nine pairs of differentially expressed metabolites and the genes ( p < 0.05) were obtained. Interestingly, a gene named TRINITY&#95;DN2608 (encoding a type of alpha-D-glucose) cloned and its overexpression lines in Arabidopsis thaliana was generated. Overexpression of TRINITY&#95;DN2608 gene elevated the susceptibility to drought stress possibly by suppressing the glucose level., Conclusion: The enrichment of sugar-related pathways advocates the potential role of glucose metabolism in drought stress. Our study provides theoretical information related to the glucose-mediated drought response and would be fruitful in future lily breeding programs., Competing Interests: The authors declare that they have no competing interests., (© 2023 Cui et al.)

Lilies ( Lilium spp.) are famous bulb flowers worldwide, with high ornamental value. Aphid damage has seriously constrained the development of the lily industry. In this study, the aphid resistance of 16 lily cultivars and 2 wild lily species was characterized in the field and greenhouse. Leaf color parameters, stomatal density and size, thickness of leaf layers, leaf waxy content, and leaf water content were determined to explore the constitutive resistance of lilies. The results show that there was a significant positive correlation between the number of aphids in the field and in the greenhouse ( p ≤ 0.05, r = 0.47). This indicated that the level of aphid infestation in both the field and the greenhouse is generally consistent across different types of lily plants. Among these 18 lilies, 'Palazzo', 'Nymph', 'Cameleon' and L. lancifolium were resistant to A. gossypii , while 'Black Beauty' and 'Magnefique' had poor resistance. The correlation analysis results showed that the number of aphids was negatively correlated with leaf abaxial surface a*, stomatal size, water content, and thickness of leaf palisade tissue and positively correlated with leaf distal axial surface b*, C*, and waxy content. Among them, the correlation between the number of aphids and the thickness of leaf palisade tissue reached a significant level ( p ≤ 0.05, r = -0.521). This indicated that the thickness of the palisade tissue of lily leaves might be an important factor influencing the proliferation of aphids. This study not only screened out aphid-resistant lilies but also established a crucial research foundation for the targeted breeding and molecular breeding of lilies with aphid resistance.

Lily is a bulbous plant with an endogenous dormancy trait. Fine-tuning bulb dormancy release is still a challenge in the development of bulb storage technology. In this study, we identified three regulators of symplastic transport, 2,3-Butanedione oxime (BDM), N-Ethyl maleimide (NEM), and 2-Deoxy-D-glucose (DDG), that also regulate bulb dormancy release. We demonstrated that BDM and DDG inhibited callose synthesis between cells and promoted symplastic transport and soluble sugars in the shoot apical meristem (SAM), eventually accelerating bulb dormancy release and flowering in lilies. Conversely, NEM had the opposite effect. These three regulators can be flexibly applied to either accelerate or delay lily bulb dormancy release., (© 2023. The Author(s).)

Petal spots are widespread in plants, they are important for attracting pollinators and as economic traits in crop breeding. However, the genetic and developmental control of petal spots has seldom been investigated. To further clarify the development of petal spots formation, we performed comparative transcriptome analysis of Lilium davidii var. unicolor and Lilium davidii petals at the full-bloom stage. In comparison with the parental species L. davidii, petals of the lily variety L. davidii var. unicolor do not have the distinct anthocyanin spots. We show that among 7846 differentially expressed genes detected, LdMYB12 was identified as a candidate gene contributing to spot formation in lily petals. The expression level of LdMYB12 in the petals of L. davidii was higher than that in L. davidii var. unicolor petals. Moreover, overexpression of LdMYB12 led to the appearance of spots on the petals of L. davidii var. unicolor, accompanied by increased expression of anthocyanin synthesis-related genes. Taken together, these results indicate that abnormal expression of LdMYB12 contributes to petal spot deficiency in L. davidii var. unicolor., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Lily is a popular flower worldwide due to its elegant appearance and pleasant fragrance. Floral volatiles of lily are predominated by monoterpenes and benzenoids. While a number of genes for monoterpene biosynthesis have been characterized, the molecular mechanism underlying floral benzenoid formation in lily remains unclear. Here, we report on the identification and characterization of a novel BAHD acyltransferase gene that contributes to the biosynthesis of two related floral scent benzoate esters, ethyl benzoate and methyl benzoate, in the scented Lilium oriental hybrid 'Siberia'. The emission of both methyl benzoate and ethyl benzoate in L. 'Siberia' was found to be tepal-specific, floral development-regulated and rhythmic. Through transcriptome profiling and bioinformatic analysis, a BAHD acyltransferase gene designated LoAAT1 was identified as the top candidate gene for the production of ethyl benzoate. In vitro enzyme assays and substrate feeding assays provide substantial evidence that LoAAT1 is responsible for the biosynthesis of ethyl benzoate. It was interesting to note that in in vitro enzyme assay, LoAAT1 can also catalyze the formation of methyl benzoate, which is typically formed by the action of benzoic acid methyltransferase (BAMT). The lack of an expressed putative BAMT gene in the flower transcriptome of L. 'Siberia', together with biochemical and expression evidence, led us to conclude that LoAAT1 is also responsible for, or at least contributes to, the biosynthesis of the floral scent compound methyl benzoate. This is the first report that a member of the plant BAHD acyltransferase family contributes to the production of both ethyl benzoate and methyl benzoate, presenting a new mechanism for the biosynthesis of benzoate esters., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision., (Copyright © 2023 Yue, Wang, Li, Liu, Yin, Huang, Zhou, Li, Yu, Chen, Yu and Fan.)

Cases of ingestion of indoor poisonous plants are relatively common among animals and lead to both acute cases of poisoning and long-term exposure to harmful substances and chronic damage to the animal's health. Plants produce a large number of secondary metabolites, which serve to protect the plant from attacks by insects, parasitic plants, fungi or, for example, during reproduction. However, these metabolites can be toxic if ingested by animals or humans. Toxicologically effective components found in plants are mainly alkaloids, glycosides, saponins, terpenes and others. This review article describes in detail the most common and popular indoor poisonous plants grown in Europe, the mechanisms of action of their active substances and clinical signs of the respective poisonings. This manuscript is supplemented with rich photographic documentation of these plants not found in similar articles, and also includes a description of the treatment of individual types of poisoning.

Owners are often not aware that some common household products, medications, and plants are toxic to dogs and cats. Six cases of toxicoses due to ingestion of xylitol, amatoxin-containing mushrooms, anticoagulant rodenticide, zinc, vitamin D, and lily are presented with typical clinical and laboratory data findings. Changes in the laboratory data are explained in the context of the mechanism of action of the toxin and the organ systems affected., Competing Interests: Disclosure The authors declare no conflicts of interest., (Copyright © 2022 Elsevier Inc. All rights reserved.)

Sugar transport and distribution plays an important role in lily bulb development and resistance to abiotic stresses. In this study, a member of the Sugar Will Eventually be Exported Transporters ( SWEET ) gene family, LoSWEET14 , from Oriental hybrid lily 'Sorbonne' was identified. LoSWEET14 encodes a protein of 278 amino acids and is capable of transporting sucrose and some types of hexoses. The transcript level of the LoSWEET14 gene was significantly increased under various stress conditions including drought, cold, salt stresses, and abscisic acid (ABA) treatment. Overexpression of LoSWEET14 in tobacco ( Nicotiana tabacum ) showed that the transgenic lines had larger leaves, accumulated more soluble sugars, and were more resistant to drought, cold, and salt stresses, while becoming more sensitive to ABA compared with wild-type lines. Promoter analysis revealed that multiple stress-related cis-acting elements were found in the promoter of LoSWEET14 . According to the distribution of cis-acting elements, different lengths of 5'-deletion fragments were constructed and the LoSWEET14-pro3 ( -540 bp ) was found to be able to drive GUS gene expression in response to abiotic stresses and ABA treatment. Furthermore, a yeast one hybrid (Y1H) assay proved that the AREB/ABF (ABRE-binding protein/ABRE-binding factor) from lilies ( LoABF2 ) could bind to the promoter of LoSWEET14 . These findings indicated that LoSWEET14 is induced by LoABF2 to participate in the ABA signaling pathway to promote soluble sugar accumulation in response to multiple abiotic stresses.

Taxonomy: Plantago asiatica mosaic virus belongs to the genus Potexvirus in the family Alphaflexiviridae of the order Tymovirales., Virion and Genome Properties: Plantago asiatica mosaic virus (PlAMV) has flexuous virions of approximately 490-530 nm in length and 10-15 nm in width. The genome of PlAMV consists of a single-stranded, positive-sense RNA of approximately 6.13 kb. It contains five open reading frames (ORFs 1-5), encoding a putative viral polymerase (RdRp), movement proteins (triple gene block proteins, TGBp1-3), and coat protein (CP), respectively., Host Range: PlAMV has an exceptionally wide host range and has been isolated from various wild plants, including Plantago asiatica, Nandina domestica, Rehmannia glutinosa, and other weed plants. Experimentally PlAMV can infect many plant species including Nicotiana benthamiana and Arabidopsis thaliana. It also infects ornamental lilies and frequently causes severe necrotic symptoms. However, host range varies depending on isolates, which show significant biological diversity within the species., Genome Diversity: PlAMV can be separated into five clades based on phylogenetic analyses; nucleotide identities are significantly low between isolates in the different clades., Transmission: PlAMV is not reported to be transmitted by biological vectors. Virions of PlAMV are quite stable and it can be transmitted efficiently by mechanical contact., Disease Symptoms: PlAMV causes red-rusted systemic necrosis in ornamental lilies, but it shows much weaker, if any, symptoms in wild plants such as P. asiatica., Control: Control of the disease caused by PlAMV is based mainly on rapid diagnosis and elimination of the infected bulbs or plants., (© 2022 The Authors. Molecular Plant Pathology published by British Society for Plant Pathology and John Wiley & Sons Ltd.)

Lilies (Lilium spp.) are one of the most important ornamental flower crops grown in Korea. Most viral diseases in lilies are transmitted by infected bulbs, which cause serious economic losses due to reduced yields. Various diagnostic techniques and high-throughput sequencing methods have been used to detect lily viruses. According to Oxford Nanopore Technologies (ONT), MinION is a compact and portable sequencing device. In this study, three plant viruses, lily mottle, lily symptomless, and plantago asiatica mosaic virus, were detected in lily samples using the ONT platform. As a result of genome assembly of reads obtained through ONT, 100% coverage and 90.3-93.4% identity were obtained. Thus, we show that the ONT platform is a promising tool for the diagnosis and characterization of viruses that infect crops.

A series polysaccharide samples extracted from three edible lilies ( Lilium davidii var. willmottiae , Lilium brownii var. viridulum , and Lilium lancifolium ) by subcritical water and ultrasound-assisted extraction were systematically compared. The results showed that extraction method was a more important factor than lily species. Subcritical water extracted lily polysaccharides (S-LP) with higher yield, molecular weight, neutral glucose and uronic acid content as well as apparent viscosity. Ultrasound-assisted extracted lily polysaccharides (U-LP) with higher reducing sugars and protein content. Moreover, due to the degradation of glycosidic bonds, ultrasonic extraction was easier to obtain lower molecular weight polysaccharides. In addition, the extraction method significantly affected the monosaccharide proportion of polysaccharides, but had no effect on type. Glucose was the main component in S-LP, and glucose and mannose were the main components in U-LP. The micromorphology of different polysaccharide samples was similar, and the scanning electron microscope (SEM) images showed regular/irregular particle clusters with different particle sizes. Overall, the relationships between extraction methods, lily species and polysaccharide properties were preliminarily elucidated, providing a reference for the targeted extraction of specific lily polysaccharides (LP)., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Song, Zhang, Luo, Ti, Wang, Ban, Tang, Hou, Xu, Ming and Yang.)

Key Message: The correlation between dormancy release and metabolic metabolic changes in lily bulbs during low temperature storage was investigated. Low temperature is a major environmental factor required for dormancy release in lily bulbs. Although great advances in plant metabolomics have been achieved, knowledge about the molecular basis of lily bulb metabolomes at different developmental stages in response to low temperature is still limited. In this work, the dormancy release, vegetative growth, flowering, metabolic profile and gene expression in the less dormant cultivar Lilium longiforum × Oriental hybrid 'Triumphator' (T) and the more dormant cultivar Lilium Asiatic hybrid 'Honesty' (H) were compared. Exposure to low temperature (LT) successfully promoted stem elongation, floral transition and flowering of both T and H bulbs. However, exposure to room temperature (RT) restricted stalk elongation of both T and H bulbs, and prohibited floral transition and flowering of H bulbs. Correspondingly, higher antioxidant enzyme activity and total primary metabolite contents were observed in the apical bud of T bulbs. Gene expression analysis revealed that expressions of LiFT, LiFLK, LiSOC1 and LiCBF were decreased, whereas the expression of LiSVP and LiFLC were increased, in the apical bud of H bulbs under RT storage condition. Our findings reveal that the growth and dormancy breaking of lily bulbs are closely associated with the metabolic changes in the apical buds during postharvest storage., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Near-infrared spectroscopy technique is a prevailing tool for quality control of foods and traditional Chinese medicines. However, it usually faced the problems of severe peak overlap, low classification accuracy and poor specificity. In this work, the potential of carbon dot-tetramethoxyporphyrin nanocomposite-based nano-effect near-infrared spectroscopy sensor combined with chemometric method was investigated for the accurate identification lily from different geographical origins. Partial least squares-discriminant analysis (PLS-DA) was used for differentiating geographical origins of lily based on the collected traditional and nano-effect near-infrared spectroscopy. Compared with traditional near-infrared spectroscopy, the nano-effect near-infrared spectroscopy obtains superior classification performance with 100% accuracy on the training and test set. The results showed that the proposed method based on near-infrared spectroscopy combined with nanocomposites and chemometrics could be considered as a promising tool for rapid discrimination of the authenticity of food and traditional Chinese medicine in the future., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

During the growth cycle of lilies, assimilates undergo a process of accumulation, consumption and reaccumulation in bulbs and are transported and allocated between aboveground and underground organs and tissues. The sink-source relationship changes with the allocation of assimilates, affecting the vegetative growth and morphological establishment of lilies. In this study, the carbohydrate contents in different tissues of five critical stages during lily development were measured to observe the assimilates allocation. The results showed bulbs acted as the main source to provide energy before the budding stage (S3); after the flowering stage (S4), bulbs began to accumulate assimilates as a sink organ again. During the period when the plant height was 30cm with leaf-spread (S2), leaves mainly accumulated assimilates from bulbs through the symplastic pathway, while when leaves were fully expanded, it transformed to export carbohydrates. At the S4 stage, flowers became a new active sink with assimilates influx. To further understand the allocation of assimilates, 16 genes related to sugar transport and metabolism ( ST genes) were identified and categorized into different subfamilies based on the phylogenetic analysis, and their protein physicochemical properties were also predicted. Tissue-specific analysis showed that most of the genes were highly expressed in stems and petals, and it was mainly the MST (monosaccharide transporter) genes that were obviously expressed in petals during the S4 stage, suggesting that they may be associated with the accumulation of carbohydrates in flowers and thus affect flower development process. LoSWEET14 (the Sugar will eventually be exported transporters) was significantly correlated with starch in scales and with soluble sugar in leaves. Sugar transporters LoHXT6 and LoSUT1 were significantly correlated with soluble sugar and sucrose in leaves, suggesting that these genes may play key roles in the accumulation and transportation of assimilates in lilies. In addition, we analyzed the expression patterns of ST genes under different abiotic stresses, and the results showed that all genes were significantly upregulated. This study lays a solid foundation for further research on molecular mechanism of sink-source change and response to abiotic stresses in lilies.

Oxalic acid (OA) is a crucial pathogenic factor for Sclerotinia spp. fungi, which is closely related to Botrytis spp. fungi. Whether OA is a pathogenic factor for the causal agent of grey mould in lily, Botrytis elliptica, and the response of lily to OA are poorly understood. To address these questions, lesion tissues and deposition of calcium oxalate (CaOX) and callose were observed in diploid and tetraploid leaves of L. rosthornii after inoculation with B. elliptica. Oxalate oxidase (OXO) activity and the transcript levels of some genes related to OA degradation (LrGLP1, LrGLP2 and LrWRKY4), reactive oxygen species (ROS) production/scavenging systems (LrRBOHD, LrGST, LrPOD and LrAPX1) and pathogen-related protein (PR) synthesis (LrCHI, LrBGL and LrPR10) were compared. After diploid and tetraploid leaves inoculation, lesion tissue and callose and CaOX were separately observed around in guard cells and stomata rather than the epidermis in the infected area. OXO activity was triggered at 2 h post-inoculation (hpi) in both ploidy leaves, and it was higher in the latter from 12-48 hpi. Expression of LrGLP1, LrGLP2, LrRBOHD, LrGST, LrPOD, LrCHI, LrBGL and LrPR10 was higher in tetraploids than in diploids from 24(12)-36(48) hpi. In conclusion, for B. elliptica, OA mainly chelates Ca 2+ from the stomata cell wall. The strong capability to degrade OA and higher expression levels of some genes related to ROS accumulation/scavenging and PR synthesis may partially explain the relatively higher grey mould resistance of tetraploid L. rosthornii., (© 2022 German Society for Plant Sciences and The Royal Botanical Society of the Netherlands.)

Lily is a popular and economically ornamental crop around the world. However, its high production of pollen grains causes serious problems to consumers, including allergies and staining of clothes. During anther development, the tapetum is a crucial step for pollen formation and microspore release. Therefore, it is important to understand the mechanism of tapetum degradation and microspore development in lily where free pollen contamination occurs. Here, we used the cut lily cultivar 'Siberia' to characterize the process of tapetum degradation through the use of cytology and transcriptomic methods. The cytological observation indicated that, as the lily buds developed from 4 cm (Lo 4 cm) to 8 cm (Lo 8 cm), the tapetum completed the degradation process and the microspores matured. Furthermore, by comparing the transcriptome profiling among three developmental stages (Lo 4 cm, Lo 6 cm and Lo 8 cm), we identified 27 differentially expressed genes. These 27 genes were classed into 4 groups by function, namely, cell division and expansion, cell-wall morphogenesis, transcription factors, LRR-RLK (leucine-rich repeat receptor-like kinases), plant hormone biosynthesis and transduction. Quantitative real-time PCR was performed as validation of the transcriptome data. These selected genes are candidate genes for the tapetum degradation and microspore development of lily and our work provides a theoretical basis for breeding new lily cultivars without pollen.

Flower color is the decisive factor that affects the commercial value of ornamental flowers. Therefore, it is important to study the regulation of flower color formation in lily to discover the positive and negative factors that regulate this important trait. In this study, MYB transcription factors (TFs) were characterized to understand the regulatory mechanism of anthocyanin biosynthesis in lily. Two R2R3-MYB TFs, LvMYB5, and LvMYB1, were found to regulate anthocyanin biosynthesis in lily flowers. LvMYB5, which has an activation motif, belongs to the SG6 MYB protein subgroup of Arabidopsis thaliana . Transient expression of LvMYB5 indicated that LvMYB5 can promote coloration in Nicotiana benthamiana leaves, and that expression of LvMYB5 increases the expression levels of NbCHS , NbDFR , and NbANS . VIGS experiments in lily petals showed that the accumulation of anthocyanins was reduced when LvMYB5 was silenced. Luciferase assays showed that LvMYB5 can promote anthocyanin synthesis by activating the ANS gene promoter. Therefore, LvMYB5 plays an important role in flower coloration in lily. In addition, the transient expression experiment provided preliminary evidence that LvMYB1 (an R2R3-MYB TF) inhibits anthocyanin synthesis in lily flowers. The discovery of activating and inhibitory factors related to anthocyanin biosynthesis in lily provides a theoretical basis for improving flower color through genetic engineering. The results of our study provide a new direction for the further study of the mechanisms of flower color formation in lilies., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Yin, Zhang, Zhang, Wang, Zhao, Irfan, Chen and Feng.)

Key Message: A number of potential genes and pathways involved in tepal trichome development were identified in a natural lily mutant by transcriptome analysis and were confirmed with trichome and trichomeless species. Trichome is a specialized structure found on the surface of the plant with an important function in survival against abiotic and biotic stress. It is also an important economic trait in crop breeding. Extensive research has investigated the foliar trichome in model plants (Arabidopsis and tomato). However, the developmental mechanism of tepal trichome remains elusive. Lilium pumilum is an edible ornamental bulb and a good breeding parent possessing cold and salt-alkali resistance. Here, we found a natural mutant of Lilium pumilum grown on a highland whose tepals are covered by trichomes. Our data indicate that trichomes of the mutant are multicellular and branchless. Notably, stomata are also developed on the tepal of the mutant as well, suggesting there may be a correlation between trichome and stomata regulation. Furthermore, we isolated 27 differentially expressed genes (DEGs) by comparing the transcriptome profiling between the natural mutant and the wild type. These 27 genes belong to 4 groups: epidermal cell cycle and division, trichome morphogenesis, stress response, and transcription factors. Quantitative real-time PCR in Lilium pumilum (natural mutant and the wild type) and other lily species (Lilium leichtlinii var. maximowiczii/trichome; Lilium davidii var. willmottiae/, trichomeless) confirmed the validation of RNA-seq data and identified several trichome-related genes., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Lily ( Lilium spp.) is an important commercial flower crop, but its market popularity and applications are adversely affected by severe pollen pollution. Many studies have examined pollen development in model plants, but few studies have been conducted on flower crops such as lily. GAMYBs are a class of R2R3-MYB transcription factors and play important roles in plant development and biotic resistance; their functions vary in different pathways, and many of them are involved in anther development. However, their function and regulatory role in lily remain unclear. Here, the GAMYB homolog LoMYB33 was isolated and identified from lily. The open reading frame of LoMYB33 was 1620 bp and encoded a protein with 539 amino acids localized in the nucleus and cytoplasm. Protein sequence alignment showed that LoMYB33 contained a conserved R2R3 domain and three BOX motifs (BOX1, BOX2, and BOX3), which were unique to the GAMYB family. LoMYB33 had transcriptional activation activity, and its transactivation domain was located within 90 amino acids of the C-terminal. LoMYB33 was highly expressed during the late stages of anther development, especially in pollen. Analysis of the promoter activity of LoMYB33 in transgenic Arabidopsis revealed that the LoMYB33 promoter was highly activated in the pollen of stage 12 to 13 flowers. Overexpression of LoMYB33 in Arabidopsis significantly retarded growth; the excess accumulation of LoMYB33 also negatively affected normal anther development, which generated fewer pollen grains and resulted in partial male sterility in transgenic plants. Silencing of LoMYB33 in lily also greatly decreased the amount of pollen. Overall, our results suggested that LoMYB33 might play an important role in the anther development and pollen formation of lily., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Liu, Wu, Feng, Yuan, He, Zhang and Teng.)

Salt stress is a major constraint of crop productivity because it reduces yield and limits the expansion of agriculture. This study investigated salt tolerance in 26 cultivars of cut lilies ( Lilium hybrids) by examining the effect of salt stress on the growth and morphological characteristics of flowers and leaves and their physiological properties (chlorophyll a fluorescence). Salt stress significantly affected the growth and development of cut lilies. Canonical discriminant analysis indicates that the middle leaf width, number of flowers, first flower diameter, petal width, and chlorophyll a fluorescence were correlated with salt stress, whereas plant height, the middle leaf length, days to flowering, and sepal width were less affected by the stress. The cultivars examined were divided into three groups: Group 1 included the salt-sensitive cultivars, which failed to develop normal flowers; Group 2 included cultivars sensitive to salt stress but tolerant to osmotic stress; and Group 3 was the salt-tolerant group, which developed commercially valuable flowers. In conclusion, the cultivars contained a variable range of cut flower characteristics and growth traits that can be employed for lily breeding programs and as material for molecular mechanisms and signaling networks under salt stress.

In plants, the cell fates of a vegetative cell (VC) and generative cell (GC) are determined after the asymmetric division of the haploid microspore. The VC exits the cell cycle and grows a pollen tube, while the GC undergoes further mitosis to produce two sperm cells for double fertilization. However, our understanding of the mechanisms underlying their fate differentiation remains limited. One major advantage of the nuclear proteome analysis is that it is the only method currently able to uncover the systemic differences between VC and GC due to GC being engulfed within the cytoplasm of VC, limiting the use of transcriptome. Here, we obtained pure preparations of the vegetative cell nuclei (VNs) and generative cell nuclei (GNs) from germinating lily pollens. Utilizing these high-purity VNs and GNs, we compared the differential nucleoproteins between them using state-of-the-art quantitative proteomic techniques. We identified 720 different amount proteins (DAPs) and grouped the results in 11 fate differentiation categories. Among them, we identified 29 transcription factors (TFs) and 10 cell fate determinants. Significant differences were found in the molecular activities of vegetative and reproductive nuclei. The TFs in VN mainly participate in pollen tube development. In comparison, the TFs in GN are mainly involved in cell differentiation and male gametogenesis. The identified novel TFs may play an important role in cell fate differentiation. Our data also indicate differences in nuclear pore complexes and epigenetic modifications: more nucleoporins synthesized in VN; more histone variants and chaperones; and structural maintenance of chromosome (SMC) proteins, chromatin remodelers, and DNA methylation-related proteins expressed in GN. The VC has active macromolecular metabolism and mRNA processing, while GC has active nucleic acid metabolism and translation. Moreover, the members of unfolded protein response (UPR) and programmed cell death accumulate in VN, and DNA damage repair is active in GN. Differences in the stress response of DAPs in VN vs. GN were also found. This study provides a further understanding of pollen cell differentiation mechanisms and also a sound basis for future studies of the molecular mechanisms behind cell fate differentiation., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 You, Zhang, Yang, Wang, Yao, Jin, Wang, Hu and Yang.)

Background: We have previously performed an analysis of the cold-responsive transcriptome in the mature leaves of tiger lily ( Lilium lancifolium ) by gene co-expression network identification. The results has revealed that a ZFHD gene, notated as encoding zinc finger homeodomain protein, may play an essential regulating role in tiger lily response to cold stress., Methods: A further investigation of the ZFHD gene (termed as LlZFHD4 ) responding to osmotic stresses, including cold, salt, water stresses, and abscisic acid (ABA) was performed in this study. Based on the transcriptome sequences, the coding region and 5' promoter region of LlZFHD4 were cloned from mature tiger lily leaves. Stress response analysis was performed under continuous 4 °C, NaCl, PEG, and ABA treatments. Functional characterization of LlZFHD4 was conducted in transgenic Arabidopsis , tobacco, and yeast., Results: LlZFHD4 encodes a nuclear-localized protein consisting of 180 amino acids. The N-terminal region of LlZFHD4 has transcriptional activation activity in yeast. The 4 °C, NaCl, PEG, and ABA treatments induced the expression of LlZFHD4 . Several stress- or hormone-responsive cis-acting regulatory elements (T-Box, Box I. and ARF) and binding sites of transcription factors (MYC, DRE and W-box) were found in the core promoter region (789 bp) of LlZFHD4 . Also, the GUS gene driven by LlZFHD4 promoter was up-regulated by cold, NaCl, water stresses, and ABA in Arabidopsis . Overexpression of LlZFHD4 improved cold and drought tolerance in transgenic Arabidopsis ; higher survival rate and better osmotic adjustment capacity were observed in LlZFHD4 transgenic plants compared to wild type (WT) plants under 4 °C and PEG conditions. However, LlZFHD4 transgenic plants were less tolerant to salinity and more hypersensitive to ABA compared to WT plants. The transcript levels of stress- and ABA-responsive genes were much more up-regulated in LlZFHD4 transgenic Arabidopsis than WT. These results indicate LlZFHD4 is involved in ABA signaling pathway and plays a crucial role in regulating the response of tiger lily to cold, salt and water stresses., Competing Interests: The authors declare that they have no competing interests., (© 2021 Yong et al.)

Description This original pen-and-ink artwork features a lily on the water symbolizing peace. As war, distrust, chaos and coups wreak havoc throughout many developing nations, healthcare workers and public health officials should be encouraged that we do in fact have a part to play in managing the human health affected by the disruption. Health work can provide a zone of comfort, spread solidarity and understanding, and manage the human toll of war. Healthcare workers have major roles in peace-keeping efforts as a result of these determinations and their impact on peace should be acknowledged and lauded. We hope that as we move forward, we can continue to improve and create positive healthcare outcomes through our public health and social work initiatives., Competing Interests: Conflicts of Interest The author declares she has no conflicts of interest., (© 2021 HCA Physician Services, Inc. d/b/a Emerald Medical Education.)

Bowman-Birk inhibitors (BBIs) are plant-derived serine proteinase inhibitors. Endogenously, they function as defense molecules against pathogens and insects, but they also have been explored for applications in cancer treatment and inflammatory disorders. Here, we isolated 15 novel BBIs from the bulb of Hyacinthus orientalis (termed HOSPIs). These isoinhibitors consisted of two or three chains, respectively, that are linked by disulfides bonds based on proposed cleavage sites in the canonical BBI reactive site loop. They strongly inhibited trypsin (Ki = 0.22-167 nM) and α-chymotrypsin (Ki = 19-1200 nM). Notably, HOSPI-B4 contains a six-residue reactive loop, which appears to be the smallest such motif discovered in BBIs to date. HOSPI-A6 and -A7 contain an unusual reactive site, i.e. Leu-Met at the P1-P1' position and have strong inhibitory activity against trypsin, α-chymotrypsin, and elastase. Analysis of the cDNA encoding HOSPIs revealed that the precursors have HOSPI-like domains repeated at least twice with a defined linker sequence connecting individual domains. Lastly, mutational analysis of HOSPIs suggested that the linker sequence does not affect the inhibitory activity, and a Thr residue at the P2 site and a Pro at the P3' site are crucial for elastase inhibition. Using mammalian proteases as representative model system, we gain novel insight into the sequence diversity and proteolytic activity of plant BBI. These results may aid the rational design of BBI peptides with potent and distinct inhibitory activity against human, pathogen, or insect serine proteinases., (© 2021 The Author(s). Published by Portland Press Limited on behalf of the Biochemical Society.)

The bulbs of several Lilium species are considered to be both functional foods and traditional medicine in northern and eastern Asia. Considering the limited information regarding the specific bioactive compounds contributing to the functional properties of these bulbs, we compared the secondary metabolites of ten Lilium bulb samples belonging to five different species, using an ultrahigh-performance liquid chromatography-electrospray ionization-tandem mass spectrometry (UPLC-ESI-MS/MS)-based secondary metabolomics approach. In total, 245 secondary metabolites were detected; further, more metabolites were detected from purple Lilium bulbs (217 compounds) than from white bulbs (123-171 compounds). Similar metabolite profiles were detected in samples within the same species irrespective of where they were collected. By combining herbal analysis and screening differential metabolites, steroid saponins were considered the key bioactive compounds in medicinal lilies. Of the 14 saponins detected, none were accumulated in the bulbs of L . davidii var. willmottiae, also called sweet lily. The purple bulbs of L . regale accumulated more secondary metabolites, and, notably, more phenolic acid compounds and flavonoids. Overall, this study elucidates the differential metabolites in lily bulbs with varying functions and colors and provides a reference for further research on functional foods and the medicinal efficacy of Lilium species.

Rhodococcus fascians is an important pathogen that infects various herbaceous perennials and reduces their economic value. In this study, we examined R. fascians isolates carrying a virulence gene from symptomatic lily plants grown in South Korea. Phylogenetic analysis using the nucleotide sequences of 16S rRNA, vicA , and fasD led to the classification of the isolates into four different strains of R. fascians . Inoculation of Nicotiana benthamiana with these isolates slowed root growth and resulted in symptoms of leafy gall. These findings elucidate the diversification of domestic pathogenic R. fascians and may lead to an accurate causal diagnosis to help reduce economic losses in the bulb market.

In bulb crops, bulbing is a key progress in micropropagation and is the feature that most distinguishes bulbous crops from other plants. Generally, bulbing involves a shoot-to-bulblet transition; however, the underlying mechanism remains elusive. We explored this process by tracking the shoot-to-bulblet transition under different culture conditions. Rapid starch accumulation occurred at 15 days after transplanting (DAT) in the bulblet-inducing treatments as confirmed via histological observations and the significant elevation of starch synthesis related-gene transcription, including LohAGPS , LohAGPL , LohGBSS , LohSS , and LohSBE . However, for shoots that did not transition to bulblets and maintained the shoot status, much higher soluble sugars were detected. Interestingly, we observed a clear shift from invertase-catalyzed to sucrose synthase-catalyzed sucrose cleavage pattern based on the differential expression of LohCWIN and LohSuSy during the key transition stage (prior to and after bulbing at 0-15 DAT). Shoots that transitioned into bulblets showed significantly higher LohSuSy expression, especially LohSuSy4 expression, than shoots that did not transition. A symplastic phloem unloading pathway at the bulblet emergence stage (15 DAT) was verified via the 6(5)-carboxyfluorescein diacetate fluorescent tracer. We propose that starch is the fundamental compound in the shoot-to-bulblet transition and that starch synthesis is likely triggered by the switch from apoplastic to symplastic sucrose unloading, which may be related to sucrose depletion. Furthermore, this study is the first to provide a complete inventory of the genes involved in starch metabolism based on our transcriptome data. Two of these genes, LohAGPS1.2b and LohSSIIId , were verified by rapid amplification of cDNA ends cloning, and these data will provide additional support for Lilium research since whole genome is currently lacking., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Wu, Ren, Gao, Sun, Li, Min, Wu, Li, Wang, Wei and Xia.)

In this study, starch extracted from lily bulbs were modified using an ultra-high pressure (UHP) treatment at six different pressure levels (100, 200, 300, 400, 500, and 600 MPa). The effects of UHP treatment on the physicochemical and morphological properties of lily starch were investigated. The morphological observation revealed that UHP treatment led to particle expansion and aggregation. Compared with the native and lily starch treated at 100-500 MPa, the lily starch treated at 600 MPa exhibited almost completely disrupted morphology and a larger particle size, indicating nearly complete gelatinization of the starch. The relative crystallinity of the UHP-treated starch remarkably reduced. Gelatinization temperatures via differential scanning calorimetry decreased with increasing pressure. The rapid viscoanalyzer results revealed that the lily starch treated with UHP at 600 MPa showed low values of peak viscosity, trough viscosity, breakdown, final viscosity, and setback. These results indicated that UHP was an effective physical modification method for lily starch, UHP treatment (600 MPa, 30 min) caused nearly complete gelatinization of lily starch, and lily starch modified using UHP might expand the application of lily in the food field., Competing Interests: The authors declare no conflict of interest., (© 2020 The Authors. Food Science & Nutrition published by Wiley Periodicals LLC.)

Rhodococcus fascians is a bacterium that causes growth abnormalities such as leafy galls, fasciation, and shoot proliferation in many plants, including ornamental plants. In February 2020, the Animal and Plant Quarantine Agency of South Korea detected 492,000 contaminated lily bulbs using an in-house PCR test based on the R. fascians fasD gene, and subsequently 1.3 million imported bulbs were destroyed. Because no pathogen isolation was associated with this diagnosis, there has been great cultivator demanded for bacterial isolation evidence of lily bulb infection with pathogenic R. fascians. To isolate the causal bacterium of the PCR tests, we sampled leaf, stem, and bulb tissues from 130 lilies with growth abnormality symptoms, collected from 24 South Korean mass production lily farms from June to August 2020. Supernatants of the homogenized samples were spread on mD2 medium (Kado and Heskett 1970) and incubated at 28°C for 10 days. Yellow to orange colonies were isolated into pure culture on mD2. Total DNA was extracted from cultures grown in yeast extract broth (YEB) at 28°C for 24 hours with Wizard DNA prep kit (Promega, Madison, WI, USA). PCR was performed to test for pathogenicity genes fas (A,D, and R) and att (A and R) (Putnam and Miller 2007). Colonies that produced at least one amplicon from these pathogenicity genes were analyzed by partial 16s rRNA gene sequencing to determine the corresponding species. Three strains that were isolated from the bulbs of fasciated lilies from Wanju (35°56´22.1˝N; 127°08´52.0˝E), Gwacheon (37°26´51.6˝N; 127°00´11.8˝E), and Yeongwol (37°18´45.8˝N; 128°11´05.6˝E), or W1, G3, and Y5 strains, yielded PCR products of the expected size for fas and att genes with the primer sets published in Serdani et al. (2013) and developed in this study (attAF: 5'-CCCGGCTACACGCATTCGC-3', attAR: 5'-CGAACGCGGTGTGCAGGT-3' and attRF: 5'-AGTGTCCCGTCGGCGAG-3', attRR: 5'-CGCGGCAGATCGAAGTCCT-3'). Sequences of the three strains were deposited in Genbank for fasA (accession MW122940-942), fasD (G3:MW122935 and 936), and fasR (MW122937-939); all shared 98.3 - 100% nucleotide identity to corresponding sequences from phytopathogenic R. fascians A25f (CP049745.1 Protein&#95;ID fasA:QII09280.1, fasD:QII09282.1, and fasR:QII09277.1). The attA and attR products were only present in G3 (attA: MW122943 and attR: MW122944) and resulted in 100% identity to those of A25f (CP049745.1 Protein&#95;ID attA:QII09269.1, attR:QII09267.1). Partial 16s rRNA gene sequences were obtained (MW064131-133) and clustered with phytopathogenic R. fascians strains D188, A21d2, and A25f. Thus we concluded that strains (W1, G3, and Y5) corresponded to R. fascians. To test the pathogenicity of these three strains, 10 seeds of garden peas for each strain were inoculated at 108 CFU/ml according to Nikolaeva et al. (2009), and the length of the main stem of each seedling was calculated 22 days post-inoculation. Seedlings inoculated with G3 and Y5 resulted in a stunted phenotype with up to 40% height reduction (p ≤ 0.001) compared to non-inoculated seedlings. As for the seedlings inoculated with W1, they exhibited as much as 15% height reduction (p ≤ 0.001). Colonies were recovered from the inoculated seedlings, identity was confirmed through colony PCR for fas and att genes. To our knowledge, this is the first report of phytopathogenic R. fascians in lilies cultivated in South Korea.

Background: Petals are the colorful region of many ornamental plants. Quality traits of petal color directly affect the value of ornamental plants. Although the regulatory mechanism of flower color has been widely studied in many plants, that of lily flower color is still worth further exploration., Results: In this study, the pigmentation regulatory network in different regions of the petal of lily cultivar 'Vivian' was analyzed through tissue structure, metabolites biosynthesis, and gene expression. We found that cell morphology of the petal in un-pigmented region differed from that in pigmented region. The cell morphology tends to flatten in un-pigmented region where the color is lighter. Moreover, high level anthocyanin was found in the pigmented regions by metabonomic analysis, especially cyanidin derivatives. However, flavanones were accumulated, contrast with anthocyanin in the un-pigmented regions of lily petal. To understand the relationship of these different metabolites and lily flower color, RNA-Seq was used to analyze the differentially expressed genes-related metabolite biosynthesis. Among these genes, the expression levels of several genes-related cyanidin derivatives biosynthesis were significantly different between the pigmented and un-pigmented regions, such as LvMYB5, LvMYB7, LvF3'H, LvDFR, LvANS and Lv3GT., Conclusions: This data will help us to further understand the regulation network of lily petal pigmentation and create different unique color species.

The Lily mottle virus (LMoV) impedes the growth and quality of lily crops in Lanzhou, China. Recently Arabis mosaic virus (ArMV) has been detected in LMoV-infected plants in this region, causing plant stunting as well as severe foliar symptoms, and likely posing a threat to lily production. Consequently, there is a need to develop simple, sensitive, and reliable detection methods for these two viruses to prevent them from spreading. Reverse transcription (RT) loop-mediated isothermal amplification (LAMP) assays have been developed to detect LMoV and ArMV using two primer pairs that match six conserved sequences of LMoV and ArMV coat proteins, respectively. RT-LAMP assay results were visually assessed in reaction tubes using green fluorescence and gel electrophoresis. Our assays successfully detected both LMoV and ArMV in lily plants without the occurrence of viral cross-reactivity from other lily viruses. Optimal conditions for LAMP reactions were 65°C and 60°C for 60 min for LMoV and ArMV, respectively. Detection sensitivity for both RT-LAMP assays was a hundredfold greater than that of our comparative RT-polymerase chain reaction assays. We have also found this relatively rapid, target specific and sensitive method can also be used for samples collected in the field and may be especially useful in regions with limited or no laboratory facilities., (© The Korean Society of Plant Pathology.)

Key Message: A MADS-domain transcription factorLoSVP, which could delay flowering through vernalization pathway, was isolated from lily. MADS-domain transcription factors play important roles in plant growth and development, especially in the transition from vegetative phase to reproductive phase. However, their functions in bulbous flowering plants are largely unknown. In this work, a SHORT VEGETATIVE PHASE (SVP) encoding genes LoSVP from oriental lily was isolated. Bioinformatic analyses demonstrated that LoSVP encodes a type II MADS-box protein containing a conserved MADS-box, as well as a conserved K-box domain. Semi-quantitative reverse transcription polymerase chain reaction (RT-PCR) revealed ubiquitous expression of LoSVP in various tissues, including petals, stamens, pistils, leaves and scales. Real-time polymerase chain reaction (PCR) analyses demonstrated that LoSVP was predominantly expressed in the early stage of developing flowers. Constitutive expression of LoSVP in Arabidopsis led to significantly delayed flowering of transgenic plants. These results suggest that LoSVP is involved in plant flowering and could be used as a potential candidate gene for the genetic regulation of flowering time in higher plants.

The lily (Lilium spp.) anther contains a lot of pollen. It is not known if lily pollen contains allergens, and therefore screening pollen allergy-related proteins and genes is necessary. The pollen development period of lily 'Siberia' was determined by microscope observation. Early mononuclear microspores and mature pollens were used as sequencing materials. The analysis of the pollen transcriptome identified differentially expressed genes (DEGs), e.g., Profilin, Phl p 7 (Polcalcin), Ole e 1, and Phl p 11, which are associated with pollen allergens. The proteome analysis positively verified a significant increase in pollen allergenic protein content. The expression levels of LoProfiilin and LoPolcalcin, annotated as allergen proteins, gradually increased in mature pollen. LoProfiilin and LoPolcalcin were cloned and their open reading frame lengths were 396 bp and 246 bp, which encoded 131 and 81 amino acids, respectively. Amino acid sequence and structure alignment indicated that the protein sequences of LoProfilin and LoPolcalcin were highly conserved. Subcellular localization analysis showed that LoProfilin protein was localized in the cell cytoplasm and nucleus. LoProfilin and LoPolcalcin were highly expressed in mature pollen at the transcriptional and protein levels. A tertiary structure prediction analysis identified LoProfilin and LoPolcalcin as potential allergens in lily pollen., Competing Interests: The authors declare that they have no conflict of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, or in the decision to publish the results.

Lilium is one of the most popular flower crops worldwide, and some species are also used as vegetables and medicines. The availability of and easy access to diverse Lilium genetic resources are essential for plant genetic improvements. Cryopreservation is currently considered as an ideal means for the long-term preservation of plant germplasm. Over the last two decades, great efforts have been exerted in studies of Lilium cryopreservation and progress has been made in the successful cryopreservation of pollen, seeds and shoot tips in Lilium . Genes that exist in Lilium, including those that regulate flower shape, color and size, and that are resistant to cold stress and diseases caused by fungi and viruses, provide a rich source of valuable genetic resources for breeding programs to create novel cultivars required by the global floriculture and ornamental markets. Successful cryopreservation of Lilium spp. is a way to preserve these valuable genes. The present study provides updated and comprehensive information about the development of techniques that have advanced Lilium cryopreservation. Further ideas are proposed to better direct future studies on Lilium cryobiotechnology., Competing Interests: Competing interestsThe authors declare that they have no competing interests., (© The Author(s) 2019.)

The biological characteristics,agronomic traits,yield traits,stress resistance,quality and photosynthetic characteristics among six lily varieties were compared in order to screen out the excellent lily varieties suitable for spread planting in Hunan province. Lilium longiflorum had the longest growth period,246 days,among these six lily varieties,while others were about 170 days. The leaves of L.longiflorum,self-selected variety,L. lancifolium and L. dauricum had higher chlorophyll content. No obvious difference was found in net photosynthetic rate,stomatal conductance,transpiration rate and intercellular CO2 concentration among all varieties. The self-selected variety had the highest theoretical and actual yield,2 543. 03,1 608. 65 kg per Mu(1 Mu≈666. 7 m2),respectively,but contents of polysaccharides and flavones in bulbs were lower. All of these six lily varieties can sowing,seedling emergence,growth,flowering,mature harvest in Hunan province. L. dauricum and L. lancifolium would be provided for edible lily. L. brownie and the self-selected variety are highly susceptible varieties. L. dauricum and L. lancifolium are suitable to plant widely in disease-prone regions,due to their strong resistance. L. brownie and L. lancifolium are preferred varieties for medicinal and food using for their good quality and higher contents of polysaccharides and flavones. L. davidii had lower theoretical and actual yield,so planting extension of it should be taken into account.

Our previous studies have indicated that a partial NAC domain protein gene is strongly up-regulated by cold stress (4 °C) in tiger lily ( Lilium lancifolium ). In this study, we cloned the full-length of this NAC gene, LlNAC2 , to further investigate the function of LlNAC2 in response to various abiotic stresses and the possible involvement in stress tolerance of the tiger lily plant. LlNAC2 was noticeably induced by cold, drought, salt stresses, and abscisic acid (ABA) treatment. Promoter analysis showed that various stress-related cis-acting regulatory elements were located in the promoter of LlNAC2 ; and the promoter was sufficient to enhance activity of GUS protein under cold, salt stresses and ABA treatment. DREB1 (dehydration-responsive binding protein1) from tiger lily (LlDREB1) was proved to be able to bind to the promoter of LlNAC2 by yeast one-hybrid (Y1H) assay. LlNAC2 was shown to physically interact with LlDREB1 and zinc finger-homeodomain ZFHD4 from the tiger lily (LlZFHD4) by bimolecular fluorescence complementation (BiFC) assay. Overexpressing LlNAC2 in Arabidopsis thaliana showed ABA hypersensitivity and enhanced tolerance to cold, drought, and salt stresses. These findings indicated LlNAC2 is involved in both DREB/CBF-COR and ABA signaling pathways to regulate stress tolerance of the tiger lily., Competing Interests: The authors declare no conflict of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, or in the decision to publish the results.

Most commercial cultivars of lily are sensitive to abiotic stresses. However, tiger lily ( Lilium lancifolium L.), one of the most widely distributed wild lilies in Asia, has strong abiotic stresses resistance. Thus, it is indispensable to identify stress-responsive candidate genes in tiger lily for the stress resistance improvement of plants. In this study, a MYB related homolog (LlMYB3) from tiger lily was functionally characterized as a positive regulator in plant stress tolerance. LlMYB3 is a nuclear protein with transcriptional activation activity at C-terminus. The expression of LlMYB3 gene was induced by multiple stress treatments. Several stress-related cis-acting regulatory elements (MYBRS, MYCRS, LTRE and DRE/CRT) were located within the promoter of LlMYB3 ; however, the promoter activity was not induced sufficiently by various stresses treatments. Overexpressing LlMYB3 in Arabidopsis thaliana L. transgenic plants showed ABA hypersensitivity and enhanced tolerance to cold, drought, and salt stresses. Furthermore, we found LlMYB3 highly co-expressed with LlCHS2 gene under cold treatment; yeast one-hybrid (Y1H) assays demonstrated LlMYB3 was able to bind to the promoter of LlCHS2 . These findings suggest that the stress-responsive LlMYB3 may be involved in anthocyanin biosynthesis pathway to regulate stress tolerance of tiger lily.

Aerial bulbils, which resemble tiny bulbs, develop from axillary buds and facilitate rapid propagation of Lilium. In most species of lily, bulbils are perpetually dormant and little is known about induction of these vegetative structures. Herein, we proposed that strigolactones (SLs) may regulate the induction of bulbils in Lilium. We tested this hypothesis by isolating and investigating the expression patterns of 2 copies of the carotenoid cleavage dioxygenase8 (CCD8) gene in lily-LoCCD8a and LoCCD8b-with regard to biosynthesis of SLs. Expression analyses revealed that LoCCD8a principally is expressed during vegetative growth, whereas LoCCD8b mainly is expressed during reproductive growth. The maximum quantity of LoCCD8a transcripts was observed in the basal plate in most developmental stages, which suggests that SLs may originate from underground parts, especially the basal plate, and move upward. The effects of treatments with indole-3-acetic acid or SL analog (GR24) on outgrowth of bulbils and expression of LoCCD8 genes suggested that SLs function downstream of auxin to inhibit the outgrowth of bulbil. The expression patterns of LoCCD8a and LoCCD8b at sprouting and bulblet weighting stages also implied that SLs may function in nutrient redistribution. Our findings are expected to promote the utilization of bulbils as vegetative propagules for commercial practice., (Copyright © 2019 Elsevier GmbH. All rights reserved.)

Background: The lily is a perennial flowering plant belonging to the genus Lilium in the family Liliaceae. Most cultivated lily plants are propagated by bulbs. Therefore, numerous lily bulbs are frequently infected by diverse viruses causing viral diseases. To date, no study has examined the viromes of plants of one type with identical genetic backgrounds collected from different geographical regions., Results: Here, we examined different viromes of the lily cultivar "Sorbonne" using 172 gigabytes of transcriptome data composed of 23 libraries from four different projects for the cultivar "Sorbonne." We identified 396 virus-associated contigs from all but one library. We identified six different viruses, including Plantago asiatica mosaic virus (PlAMV), Cucumber mosaic virus (CMV), Lily symptomless virus (LSV), Tulip virus X (TVX), Lily mottle virus (LMoV), and Tobacco rattle virus (TRV). Of them, PlAMV was the most common virus infecting the lily. Scale and flower samples possessed a high number of virus-associated reads. We assembled 32 nearly complete genomes for the six identified viruses possessing the polyadenylate tails. Genomes of all six viruses were highly conserved in the lily cultivar "Sorbonne" based on mutation analysis. We identified defective RNAs from LSV, TVX, and PlAMV localized in the triple gene block region. Phylogenetic analyses showed that virus genomes are highly correlated with geographical regions and host plants., Conclusions: We conducted comprehensive virome analyses of a single lily cultivar, "Sorbonne," using transcriptome data. Our results shed light on an array of lily virome-associated topics, including virus identification, the dominant virus, virus accumulation in different plant tissues, virus genome assembly, virus mutation, identification of defective RNAs, and phylogenetic relationships of identified viruses. Taken together, we provide very useful methods and valuable results that can be applied in other virome-associated studies.