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Objective: Lilium brownii var. viridulum (LB) and L. lancifolium (LL) are the main sources of medicinal lily (Lilii Bulbus, Baihe in Chinese) in China. However, the functional components of these two species responsible for the treatment efficacy are yet not clear. In order to explore the therapeutic material basis of Lilii Bulbus, we selected L. davidii var. willmottiae (LD) only used for food as the control group to analyze the differences between LD and the other two (LB and LL). Methods: Metabolome and transcriptome were carried out to investigate the differences of active components in LD vs LB and LD vs LL. Data of metabolome and transcriptome was analysed using various analysis methods, such as principal component analysis (PCA), hierarchical cluster analysis (HCA), and so on. Differentially expressed genes (DEGs) were enriched through KEGG and GO enrichment analysis. Results: The PCA and HCA of the metabolome indicated the metabolites were clearly separated and varied greatly in LL and LB contrasted with LD. There were 318 significantly differential metabolites (SDMs) in LD vs LB group and 298 SDMs in LD vs LL group. Compared with LD group, the significant up-regulation of steroidal saponins and steroidal alkaloids were detected both in LB and LL groups, especially in LB group. The HCA of transcriptome indicated that there was significant difference in LB vs LD group, while the difference between LL and LD varied slightly. Additionally, 47 540 DEGs in LD vs LB group and 18 958 DEGs in LD vs LL group were identified. Notably, CYP450s involving in the biosynthesis of steroidal saponins and steroidal alkaloids were detected, and comparing with LD, CYP724, CYP710A, and CYP734A1 in LB and CYP90B in LL were all up-regulated. Conclusion: This study suggested that steroidal saponins and steroidal alkaloids maybe the representative functional components of Lilii Bulbus, which can provide new insights for Lilii Bulbus used in the research and development of classic famous formula.

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Lilium lancifolium Thunb (L. lancifolium) is an important medicinal and edible plant with outstanding functionality for selenium (Se) biofortification. However, the molecular response of L. lancifolium to exogenous Se has not been fully elucidated. In this study, the effects of different levels of Se on L. lancifolium growth and quality were explored by transcriptome, metabolome and biochemical analyses. The results showed that the total Se and organic Se content in L. lancifolium bulbs increased with increasing Se dosage (0-8.0 mmol/L). Moreover, Se stimulated the growth of L. lancifolium at low level (2.0 mmol/L) but showed an inhibitory effect at high levels (≥4.0 mmol/L). Metabolomic and biochemical analyses revealed that the bulb weight and the content of amino acid, soluble sugar, and soluble protein were significantly increased in the 2.0 mmol/L Se treatment compared with those in the control (0 mmol/L Se). Transcriptome and metabolome analyses revealed that the significant upregulation of the GPD1, GPAT and ADPRM genes promoted glycerophospholipid accumulation. Additionally, the significantly upregulated glyA and downregulated asnB, nadB, thrA and SAT genes coordinate to the regulation of amino acid biosynthesis. The significantly upregulated SUS, bgl B, BAM, and SGA1 genes were involved in soluble sugar accumulation under Se treatment. In summary, this study identified the optimal Se concentration (2.0 mmol/L), which significantly improved the growth and nutritional quality of L. lancifolium and contributed to understanding the combined effects of Se treatment on the expression of genes and the accumulation of metabolites in L. lancifolium bulbs.

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Four new steroidal glycosides (1-4), including two steroidal saponins named lililancifoloside B and C (1-2), one pregnane glycoside named lililancifoloside D (3), and one C22-steroidal lactone glycoside named lililancifoloside E (4), together with five known ones (5-9), were isolated from the bulbs of Lilium lancifolium Thunb. By using spectroscopic analysis, including 1D, 2D NMR, and HR-ESI-MS, the structures of 1-4 were elucidated. All isolates were tested for their cytotoxic potential against the MCF-7, MDA-MB-231, HepG2, and A549 cell lines. Compound 6 distinguished out among them, IC50 values of 3.31, 5.23, 1.78, and 1.49 µM against the four cell lines, respectively. Other compounds such as compound 3, 5, and 9 have also shown specific cytotoxic activity. Next, studies showed that compound 6 might cause HepG2 cells to undergo a cell cycle arrest during the G2/M phase and apoptosis.

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Hyperpigmentation disorders causing emotional distress require the topical use of depigmenting agents of natural origin. In this study, the anti-melanogenic effects of the Lilium lancifolium root extract (LRE) were investigated in B16F10 cells. Consequently, a non-cytotoxic concentration of the extract reduced intracellular melanin content and tyrosinase activity in a dose-dependent manner, correlating with the diminished expression of core melanogenic enzymes within cells. LRE treatment also inhibited cyclic adenosine monophosphate (cAMP) response element-binding protein (CREB)/microphthalmia-associated transcription factor signaling, which regulates the expression of tyrosinase-related genes. Upon examining these findings from a molecular mechanism perspective, LRE treatment suppressed the phosphorylation of protein kinase A (PKA), p38, and extracellular signal-related kinase (ERK), which are upstream regulators of CREB. In addition, L-phenylalanine and regaloside A, specifically identified within the LRE using liquid chromatography-mass spectrometry, exhibited inhibitory effects on melanin production. Collectively, these results imply that LRE potentially suppresses cAMP-mediated melanogenesis by downregulating PKA/CREB and mitogen-activated protein kinase (MAPK)/CREB signaling pathways. Therefore, it can be employed as a novel therapeutic ingredient of natural origin to ameliorate hyperpigmentation disorders.

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Objective @#To identify the main components in the extracts of different parts of Juandan Baihe (Lilium lancifolium) by ultra-high performance liquid chromatography quadrupole time-of-flight mass spectrometry (UPLC-Q-TOF-MS) technology and investigate their hypoglycemic activities.@*Methods@#The MS fragmentation pathways of the main types of compounds in Juandan Baihe (Lilium lancifolium) were studied, and the main components in the extracts were systematically identified using MS fragmentation pathways combined with MS mining technology. Based on the hyperglycemia male mouse model [specific pathogen free (SPF)-grade Kunming mice] induced by streptozotocin (intragastric administration of 80 mg/kg for 3 d), the hypoglycemic effects of extracts of Juandan Baihe (Lilium lancifolium) roots, stems, corms, leaves, and flowers were evaluated by measuring the changes of blood glucose, daily water consumption, daily food intake, and body weight.@*Result@#The MS fragmentation pathways of regalosides, dioscins, phenylpropanoids, flavonoids, and chlorogenic acids in Juandan Baihe (Lilium lancifolium) were clarified, and a mining method for compounds in this plant was constructed. A total of 58 compounds, including 6 chlorogenic acids, 14 regalosides, 13 phenylpropanoids, 5 flavonoids, and 20 dioscins, were identified from the roots, stems, corms, leaves, and flowers of Juandan Baihe (Lilium lancifolium). Among them, 30 compounds were reported for the first time from this plant. The root and corm extracts demonstrated significant hypoglycemic activities by reducing blood glucose levels from 23.76 ± 1.21 and 24.29 ± 1.35 mmol/L to 17.21 ± 1.23 and 18.78 ± 1.49 mmol/L, respectively (P < 0.05). The roots and corms extracts could also attenuate the symptoms of polydipsia (P < 0.01), polyphagia (P < 0.05), and weight loss caused by diabetes.@*Conclusion@#This study clarifies that the roots of Juandan Baihe (Lilium lancifolium) are rich in regalosides and dioscins for the first time, and have significant hypoglycemic activities, providing the foundation for the comprehensive utilization of this plant and the development of hypoglycemic drugs.

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Background and Aim: Osteoarthritis (OA) is a chronic, painful, degenerative inflammatory disease of the synovial joints. Regular use of nonsteroidal anti-inflammatory drugs to decrease OA pain can have severe side effects, such as gastric irritation, ulcers, and heart problems. Natural products are extensively used to minimize OA-associated pain and inflammatory reactions. Lilium lancifolium is commonly used to alleviate several diseases through its anti-inflammatory effects. This study examined the impact of L. lancifolium extract on alleviating pain and inflammation associated with articular cartilage damage. Materials and Methods: Hydro-ethanol extracts of the L. lancifolium bulb were used. The experimental animals (adult beagle dogs) were divided into four groups: sham, which received neither treatment nor surgery; placebo, which received an empty gelatin capsule; glucosamine, which received glutamine (60 mg/kg); and L. lancifolium, which received an L. lancifolium extract-filled (60 mg/kg) gelatin capsule for 8 weeks. OA was induced by an expert orthopedic surgeon in 2-year-old dogs through resection of cranial cruciate ligament and lateral collateral ligament. Inflammatory cytokines, enzymes, lameness score, radiology, and histological changes were assessed. Results: Our experiments showed that long-term oral therapy with L. lancifolium alleviated inflammation and increased histological damage. L. lancifolium treatment effectively reduced cytokines, such as interleukin-6, metalloproteinase-9, leukotriene-4, prostaglandin, and cyclo-oxygenase in dogs with OA, suggesting the potential to minimize inflammatory reactions in OA. L. lancifolium showed anti-inflammatory qualities in dogs with OA. This effect was comparable with that of glucosamine OA treatment. Conclusion: L. lancifolium supplementation represents a possible therapeutic and management option in this model of OA.

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Lilium, a perennial crop with great ornamental, medicinal and edible value, has been frequently used as functional food and medicine. Lilium lancifolium Thunb. (L. lancifolium) and Lilium brownii F.E.Brown var.viridulum Baker (L. brownii) are the most used medicinal species in China. However, the flavor compounds of these two species have not yet been clear. Here, metabolomics and transcriptome analysis were used to reveal the difference of the bitter substances of L. lancifolium and L. brownii. Qualitative results indicated that nine compounds are commonly existed in L. lancifolium and L. brownii, while nine compounds are unique in L. lancifolium and eight compounds are unique in L. brownii. Furthermore, quantitative results revealed that the content of regaloside A in L. lancifolium was nearly 2-7 folds higher than that of L. brownii, and the content of regaloside B in L. lancifolium was about 4-16 folds higher than that of L. brownii. Regaloside C and E were not detected in L. brownii. Transcriptome analysis showed that there were 90 unique genes up-regulated in L. lancifolium samples in the pathway of phenylpropanoid biosynthesis and 75 unique genes up-regulated in L. brownii samples, which could be related to the different content and chemical structure specificity of phenylpropanoid glycerol glucosides in L. lancifolium and L. brownii. The results of our in-deep research provide new insights into the bitter substances of L. lancifolium and L. brownii, and a further consideration for the chemical consistency and quality evaluation for Lilii bulbus.

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Yishui Lily 140 (Lilium lancifolium) is a hybrid lily species which was bred from wild lily varieties due to its edible and medicinal value. In this study, we have sequenced the complete chloroplast (cp) of L. lancifolium. The complete cp sequence is 152,643 bp long, with a large single copy (LSC) region of 82,084 bp, a small single copy (SSC) region of 17,513 bp, and two inverted repeat (IR) regions of 26,492 bp each. The GC contents of the complete cp genomes are 37.0%. It contains 132 genes, including 86 coding genes, 8 ribosomal RNAs, and 38 transfer RNAs. Among them, 16 different genes have a single intron and the remaining two genes have double introns, including nine cis-splicing and one trans-splicing genes. Compared with other species, we found three high variation hot spots and 96 repeats sequence. The genetic information of Lilium can be enriched as well as identifying proximal species. They are edible and have medicinal value for humans. Therefore, sequencing of Yishui Lily 140 is important to explore the cp genome composition.

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The bulbil is an important vegetative reproductive organ in triploid Lilium lancifolium whose development is promoted by cytokinins. Type-B response regulators (RRs) are critical regulators that mediate primary cytokinin responses and promote cytokinin-induced gene expression. However, the function of cytokinin type-B Arabidopsis RRs (ARRs) in regulating bulbil formation is unclear. In this study, we identified five type-B LlRRs, LlRR1, LlRR2, LlRR10, LlRR11 and LlRR12, in L. lancifolium for the first time. The five LlRRs encode proteins of 715, 675, 573, 582 and 647 amino acids. All of the regulators belong to the B-I subfamily, whose members typically contain a conserved CheY-homologous receiver (REC) domain and an Myb DNA-binding (MYB) domain at the N-terminus. As transcription factors, all five type-B LlRRs localize at the nucleus and are widely expressed in plant tissues, especially during axillary meristem (AM) formation. Functional analysis showed that type-B LlRRs are involved in bulbil formation in a functionally redundant manner and can activate LlRR9 expression. In summary, our study elucidates the process by which cytokinins regulate bulbil initiation in L. lancifolium through type-B LlRRs and lays a foundation for research on the molecular mechanism of bulbil formation in the lily.

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The Lilium lancifolium Thunb. is a herb with multiple functions in both medicine and food in China, and its extracts have shown antidepressant effects. In this study, fresh bulbs of Lilium lancifolium Thunb. were processed to study the effects of different drying processes on changes in its main chemical components. We found that different drying methods can affect the chemical constituents of the herb. Among these components, Regaloside A has been found as the characteristic component. Here, Cell Counting Kit-8 assay, and Western blotting were used to evaluate the neuroprotective antidepressant effects of Regaloside A. The results showed the cell survival rate was improved, the phosphorylation levels of brain-derived neurotrophic factor, tyrosine kinase receptor B, phosphatidylinositol 3 kinase, protein kinase B, and mammalian target of rapamycin were increased after Regaloside A treatment. In general, different drying methods have a significant influence on the chemical composition of the herb, and Regaloside A may be the main chemical component of the herb. It can alleviate the damage of corticosterone in SH-SY5Y cells, and phosphatidylinositol-3-kinase/protein kinase B/mammalian target of rapamycin signaling mediated by brain-derived neurotrophic factor/tyrosine kinase receptor B may play an important role in the neuroprotective antidepressant effects of Regaloside A.

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Bulbus Lilii, as a medicinal and edible plant, has anti-inflammatory, anti-oxidative and immunopotentiating pharmacological activities, which seems to be therapeutic on cancer prevention. The purpose of this study was to investigate the effects of total saponins from Lilium lancifolium (TSLL) on proliferation, apoptosis and migration of human gastric carcinoma cells lines SGC-7901 and HGC-27 and its underlying mechanism. The results showed that TSLL inhibited the proliferation of gastric carcinoma cells by suppressing the level of proliferating cell nuclear antigen (PCNA) and increased p21 level. TSLL induced cells apoptosis by up-regulating expression of pro-apoptotic protein Bax and down-regulating anti-apoptotic protein Bcl-2 expression. Meanwhile, TSLL remarkably inhibited cell migration and invasion, decreased matrix metalloproteinase-2 (MMP-2) expression and increased tissue inhibitor of metalloproteinases-1 (TIMP-1) expression. Notably, TSLL had stronger anti-cancer effect on undifferentiated HGC-27 cells than differentiated SGC-7901 cells. Accordingly, TSLL might be a promising candidate to prevent and suppress the growth of gastric carcinoma cells.

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Although anthocyanin color patterns on flowers are among the most attractive characteristics, the genetic mechanisms through which color patterns are developed are not well understood, especially for color patterns associated with altered petal structure. Lilium species and cultivars often develop raised spots, where the interior surfaces of tepals increase to develop bumps with accompanying anthocyanin accumulation. The aim of this study was to identify transcription factors regulating pigmentation of the bumps. We identified two R2R3-MYB genes, MYB19Long and MYB19Short, in Lilium leichtlinii, L. lancifolium, and Asiatic hybrid lily cultivars. Their amino acid sequences were similar; however, part of the C-terminal region was triplicated in MYB19Long. Spatial and temporal expression profiles in lilies were strongly associated with anthocyanin biosynthesis gene expression in the bumps, and some defects were found in these genes in L. lancifolium 'Pure Gold' that developed colorless bumps. Thus, both MYB19Long and MYB19Short were likely to be involved in the bump pigmentation. MYB19Long had a stronger ability to stimulate target gene expression than MYB19Short, and expression levels of MYB19Long were greater than those of MYB19Short in lily tepals; thus, the ability to biosynthesize anthocyanin pigments was greater for MYB19Long than for MYB19Short. Among the F1 population, MYB19Short expression was found only in the tepals of F1 plants that developed bumps, although all of the F1 plants possessed the MYB19Short gene, indicating that MYB19 expression followed bump development. These findings helped to elucidate the genetic mechanisms underlying raised spot development.

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KEY MESSAGE: The cytokinin pathway promotes the initiation of bulbil formation, and iPA may an important type of cytokinin during bulbil formation in Lilium lancifolium. Bulbils are important vegetative reproductive organs in triploid Lilium lancifolium. We previously showed that cytokinins are involved in bulbil formation, but how cytokinins participate in bulbil formation is not clear. In this study, bulbil formation was divided into three stages on the basis of anatomical and histological observations: the bulbil initiation stage, bulbil primordium-formation stage and bulbil structure-formation stage. The results indicated that iPA was the most critical cytokinin during the bulbil initiation. qRT-PCR revealed that increased iPA content during bulbil initiation was mainly due to increased expression of cytokinin synthesis genes (IPT1/5) and cytokinin activation genes (LOG1/3/5/7) and significantly decreased expression of the cytokinin degradation gene CKX4. Exogenous 6-BA and lovastatin affected the cytokinin pathway and promoted or inhibited bulbil initiation by increasing or decreasing the content of endogenous iPA, respectively. In summary, we demonstrate that cytokinins positively regulate bulbil formation and provide preliminary insight into the regulatory mechanisms by which the cytokinin pathway promotes bulbil initiation.

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Lilium lancifolium is native to Northeast Asia and its bulbs have been used for medicinal treatment. Moreover, Japan has been using L. lancifolium bulbs more actively as food ingredients than Korea. Therefore, this study was to investigate the characteristics of Korean L. lancifolium bulbs, with respect to food component and functionality. As a result of proximate composition analysis, L. lancifolium bulbs have an abundant carbohydrate content. HPLC analysis indicated p-coumaric acid and ferulic acid contents of Korean L. lancifolium extract were 1.14 ± 0.01, 1.46 ± 0.00 mg/g, but only p-coumaric acid was less detected in Japanese extract. Also, Korean L. lancifolium bulbs extract exhibited significant antioxidant effects, as evaluated with antioxidant activity and compound, than Japanese extract. Furthermore, Korean L. lancifolium bulbs extract significantly inhibited pro-inflammatory protein expressions through MyD88 dependent pathway. Therefore, these results suggested Korean L. lancifolium bulbs have the potential to being functional food ingredients. PRACTICAL APPLICATIONS: Lilium lancifolium is a perennial plant belonging to the Liliaceae family. The storage organ of L. lancifolium is surrounded by several fleshy nodes at the base of the stem, called the bulb, which has been used as food or medicine to treat pneumonia and bronchitis. L. lancifolium is widely found in countries of Northeast Asia, such as Korea, Japan, and China, and its bulbs have been studied for presence of bioactive compounds that have important functional activities. The bioactive compounds in the L. lancifolium bulbs may vary from region to region. In this study, the difference observed in the contents of different bioactive compounds and the efficacy of anti-inflammatory effects of L. lancifolium bulbs from different regions were consistent in this regard. As a comparative study of food materials by region, these L. lancifolium bulbs have the potential to be used as a food material for preventing inflammatory diseases.

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Objective: To develop the EST-SSR molecular identification system of Lilium lancifolium, Lilium davidii var. willmottiae, Lilium regale, Lilium casa blanca and Lilium brownie var. viridulum, and analyze the development efficiency and identification ability of EST-SSR molecular marker technology for Lilium genus. Methods: The MISA.pl program was used to identify the SSR locus of the EST gene sequence published by NCBI. The EST-SSR primers were generated by Primer3 program module, and the primers were screened by PCR amplification and agarose gel electrophoresis. The primary screening primers were verified by polyacrylamide gel electrophoresis, and the characteristic bands of different germplasms were labeled and analyzed to construct an identification system. Results: A total of 199 pairs of SSR primers were designed. After screening 26 pairs of primers, two pairs of highly efficient primers were obtained. The molecular identification system constructed by primer JZ391 can effectively identify the mixed commercial materials, which had certain practical value. Conclusion: Based on the extreme genetic characteristics of the research materials, this identification marker development is very efficient. The results confirm that the genetic basis of the species is an important factor affecting the development efficiency of its EST-SSR molecular marker. At the same time, this case can be used as a reference for the development of EST-SSR markers for other Chinese medicinal herbs similar to Lilium genus.

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Objective To analyze and discuss the content determination of As and Hg from Lilium Lancifolium in Longshan County; To optimize microwave digestion conditions. Methods Automatic microwave digestion appratus was used. Lilium Lancifolium samples from Longshan County were digested in teflon microwave tube with HNO3-H2O2, and As and Hg were measured with atomic fluorescence spectrometry (AFS). Results The content of As was in the range of 0.031–0.507 mg/kg, and the highest content of Hg was 0.024 mg/kg. The regression equation was Y=221.23X+170.72(r=0.995 9),Y=503.52X-682.43,(r=0.999 2).For the production base,the recoveries of As and Hg were 94.32% and 92.48% in the samples, and RSD were 2.14% and 2.70%; for the breeding base, the recoveries of As and Hg were 94.95% and 93.52% in the samples, and RSD were 1.15% and 1.97%. Conclusion The method is simple and reliable, which can be used to the content determination of As and Hg from Lilium Lancifolium, and provide references for the choice of base of production and breeding of Lilium Lancifolium in Longshan County.

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Lilium lancifolium Thunb. is a Chinese traditional plant with various health benefits. In this study, we purified and characterized the water-soluble polysaccharide fraction (LLP-1A) of L. lancifolium. We also investigated the in vitro immune-enhancing activity of LLP-1A in macrophages and the underlying molecular mechanism. Results showed that LLP-1A was mainly composed of mannose and glucose at a molar ratio of 1.77:1, and its molecular weight was approximately 78.61kDa. The Fourier transform-infrared spectra of LLP-1A also revealed typical polysaccharide characteristics: the presence of uronic acid, pyranose rings and ß-glycosidic bonds. With regard to its effects on macrophages, LLP-1A enhanced phagocytic activity and induced the NO production in a dose-dependent manner. Further, it induced expression of the cytokines interleukin-6, monocyte chemotactic protein 1, tumor necrosis factor-α and interleukin-1ß. With regard to the molecular mechanism, LLP-1A increased protein expression of Toll-like receptor 4 and phosphorylation of the inhibitor of nuclear factor kappa-B kinase, inhibitor of NF-κB, and nuclear factor-kappa B in RAW 264.7 cells. Therefore, the data suggest that LLP-1A significantly upregulated the expression of immune reactive cytokines in RAW 264.7 macrophages through the TLR4-mediated NF-κB signal pathway. Thus, LLP-1A may have immunomodulatory functions that may prove beneficial for the treatment of immune-related diseases.

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This study established a rapid UPLC-TQ-MS/MS method for determination of eight active ingredients in Lilium lancifolium. The contents range of regaloside E, F, C and B are as follows: 0.604 0×10⁻¹-18.62×10⁻¹, 0.680 0×10⁻²-44.75×10⁻², 0.700 0×10⁻³-29.65×10⁻¹, 0.170 0×10⁻¹-4.724 mg•g⁻¹; the contents of chlorogenic acid, caffeic acid, protocatechualdehyde and ferulic acid, within the range of 6.827×10⁻³-16.07×10⁻³, 0.011 1×10⁻³-79.71×10⁻³, 0.593 7×10⁻³-2.962×10⁻³, 2.606×10⁻²-45.89×10⁻² mg•g⁻¹, respectively. According to PCA (principal components analysis) plotting, 35 batches can be divided into two categories, namely Anhui Huoshan and Hunan Longshan. The main different elements between these two categories are caffeic acid and ferulic acid according to the VIP (variable importance in the projection) points figure. Based on comprehensive principal component values, there are eight batches of L. lancifolium from Huoshan among the comprehensive ranking of ten. The UPLC-TQ-MS method for simultaneous analysis of eight active ingredients is accurate, efficient and convenient. This result can provide scientific basis for quality control of L. lancifolium.

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Aerial bulbils are an important propagative organ, playing an important role in population expansion. However, the detailed gene regulatory patterns and molecular mechanism underlying bulbil formation remain unclear. Triploid Lilium lancifolium, which develops many aerial bulbils on the leaf axils of middle-upper stem, is a useful species for investigating bulbil formation. To investigate the mechanism of bulbil formation in triploid L. lancifolium, we performed histological and transcriptomic analyses using samples of leaf axils located in the upper and lower stem of triploid L. lancifolium during bulbil formation. Histological results indicated that the bulbils of triploid L. lancifolium are derived from axillary meristems that initiate de novo from cells on the adaxial side of the petiole base. Transcriptomic analysis generated ~650 million high-quality reads and 11,871 differentially expressed genes (DEGs). Functional analysis showed that the DEGs were significantly enriched in starch and sucrose metabolism and plant hormone signal transduction. Starch synthesis and accumulation likely promoted the initiation of upper bulbils in triploid L. lancifolium. Hormone-associated pathways exhibited distinct patterns of change in each sample. Auxin likely promoted the initiation of bulbils and then inhibited further bulbil formation. High biosynthesis and low degradation of cytokinin might have led to bulbil formation in the upper leaf axil. The present study achieved a global transcriptomic analysis focused on gene expression changes and pathways' enrichment during upper bulbil formation in triploid L. lancifolium, laying a solid foundation for future molecular studies on bulbil formation.

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This study established a rapid UPLC-TQ-MS/MS method for determination of eight active ingredients in Lilium lancifolium. The contents range of regaloside E, F, C and B are as follows: 0.604 0×10⁻¹-18.62×10⁻¹, 0.680 0×10⁻²-44.75×10⁻², 0.700 0×10⁻³-29.65×10⁻¹, 0.170 0×10⁻¹-4.724 mg•g⁻¹; the contents of chlorogenic acid, caffeic acid, protocatechualdehyde and ferulic acid, within the range of 6.827×10⁻³-16.07×10⁻³, 0.011 1×10⁻³-79.71×10⁻³, 0.593 7×10⁻³-2.962×10⁻³, 2.606×10⁻²-45.89×10⁻² mg•g⁻¹, respectively. According to PCA (principal components analysis) plotting, 35 batches can be divided into two categories, namely Anhui Huoshan and Hunan Longshan. The main different elements between these two categories are caffeic acid and ferulic acid according to the VIP (variable importance in the projection) points figure. Based on comprehensive principal component values, there are eight batches of L. lancifolium from Huoshan among the comprehensive ranking of ten. The UPLC-TQ-MS method for simultaneous analysis of eight active ingredients is accurate, efficient and convenient. This result can provide scientific basis for quality control of L. lancifolium.