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Ultraviolet-B (UV-B) radiation is a major abiotic stress that dragon trees are exposed to during their growth and development; however, it is also an environmental signal perceived by plants that affects the flavonoid pathway. Previous studies have demonstrated that amounts of flavonoids are contained in dragon tree resin, otherwise known as dragon's blood. However, the traits and mechanism involved in the UV-B-mediated increase in flavonoids in dragon trees are still unknown. Here, we studied the response of Dracaena cambodiana under full solar UV-B radiation. The results showed that the contents of total flavonoids in D. cambodiana significantly increased after UV-B radiation exposure. Then, the transcriptome was used for determining the interactive mechanism of flavonoid accumulation and UV-B stress. Differential expression analyses identified 34 differentially expressed genes (DEGs) involved in flavonoid synthesis; specifically, 24 of the identified DEGs were significantly up-regulated after UV-B radiation exposure. In addition, 57 DEGs involved in Ca2+/kinase sensors, 58 DEGs involved in ROS scavenging and the plant hormone pathway, and 116 DEGs transcription factors in 5 families were further identified and analyzed. Finally, we deduced the potential mechanism of UV-B-promoting flavonoid formation to neutralize ROS damage derived from UV-B radiation in D. cambodiana based on the gene co-expression network and previous studies from other plants. Considering that wild dragon tree populations are currently highly threatened by anthropogenic and natural stressors, the interactive studies between D. cambodiana plants and UV-B radiation provide valuable information toward understanding the mechanism of dragon's blood formation and help us reveal the evolution of D. cambodiana, with the eventual goal of aiding in the global conservation of this precious biological resource. [ABSTRACT FROM AUTHOR]

Background: Flavonoids, found in common vegetables and fruits, have health benefits that are often limited by their low bioavailability. Excipient emulsions provide an effective strategy to overcome these obstacles. However, the nature of the emulsifier used to formulate excipient emulsions and the chemical structure of the flavonoids both affect the bioaccessibility of the flavonoids., Results: The purpose of this study was to investigate the impact of the interfacial properties of excipient emulsions on the in vitro gastrointestinal fate of representative structural flavonoids (quercetin, kaempferol, and apigenin) through the INFOGEST method. Tween 80 (TW80) (a nonionic surfactant) was more effective at reducing the oil-water interfacial tension than whey protein isolate (WPI) (a protein-based emulsifier) or octenyl succinic anhydride (OSA)-modified starch (MS) (a polysaccharide-based emulsifier). Moreover, TW80 created excipient emulsions with smaller oil droplets, which were more resistant to oral and gastric conditions. The WPI-emulsions underwent severe flocculation in the gastric phase, leading to an appreciable increase in particle size (from 220 to 3000 nm). The TW80-coated oil droplets were more digestible than WPI- or MS-coated ones. This was attributed to the larger lipid surface area for lipase attachment. The bioaccessibility of quercetin, kaempferol, and apigenin was also affected by emulsifiers: TW 80 (25% to 45%) > WPI (14% to 29%) ≈ MS (15% to 25%). Flavonoid bioaccessibility appeared to be related to their molecular properties., Conclusion: This study provides guidance for the design of effective excipient emulsions to enhance the bioavailability of flavonoids. © 2024 Society of Chemical Industry., (© 2024 Society of Chemical Industry.)

Astragali radix (HQ) is a herb with rich medicinal and edible value. Wild-simulated HQ (FYS) and Transplanted HQ (PZ) are its currently two primary forms available in the market. Metabolomics was employed to investigate their intricate metabolic variations under various cultivation methods and growth years. Notable similarities were observed in their metabolic changes across various growth years. Specifically, saponins was higher in the early growth phase, while flavonoids increased in the later. Additionally, comparative analysis of HQ samples from different cultivation methods indicated that FYS generally exhibited different chemical characteristics compared to PZ within the same market circulation period, and Calycosin-7-O-Glc-6"-O-acetate and Cycloastragenol-H 2 O might be used to discriminant them (the content of Calycosin-7-O-Glc-6"-O-acetate and Cycloastragenol-H 2 O was higher in FYS than in PZ). This approach elucidates the dynamic change pattern of characteristic metabolites and pinpoints potential biomarkers for both FYS and PZ, thereby enhancing our understanding of these medicinal materials., 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 © 2024 Elsevier Ltd. All rights reserved.)

Common bean (Phaseolus vulgaris L.) market classes have distinct seed coat colors, which are directly related to the diverse flavonoids found in the mature seed coat. To understand and elucidate the molecular mechanisms underlying the regulation of seed coat color, RNA-Seq data was collected from the black bean 5-593 and used for a differential gene expression and enrichment analysis from four different seed coat color development stages. 5-593 carries dominant alleles for 10 of the 11 major genes that control seed coat color and expression and has historically been used to develop introgression lines used for seed coat genetic analysis. Pairwise comparison among the four stages identified 6,294 differentially expressed genes (DEGs) varying from 508 to 5,780 DEGs depending on the compared stages. Kyoto Encyclopedia of Genes and Genomes (KEGG) enrichment analysis revealed that phenylpropanoid biosynthesis, flavonoid biosynthesis, and plant hormone signal transduction comprised the principal pathways expressed during bean seed coat pigment development. Transcriptome analysis suggested that most structural genes for flavonoid biosynthesis and some potential regulatory genes were significantly differentially expressed. Further studies detected 29 DEGs as important candidate genes governing the key enzymatic flavonoid biosynthetic pathways for common bean seed coat color development. Additionally, four gene models, Pv5-593.02G016100, 593.02G078700, Pv5-593.02G090900, and Pv5-593.06G121300, encode MYB-like transcription factor family protein were identified as strong candidate regulatory genes in anthocyanin biosynthesis which could regulate the expression levels of some important structural genes in flavonoid biosynthesis pathway. These findings provide a framework to draw new insights into the molecular networks underlying common bean seed coat pigment development., Competing Interests: Declarations. Ethics approval and consent to participate: Not applicable. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Citrus is an important dietary source of flavonoid glycosides, and UDP-glycosyltransferases (UGTs) are the key enzymes responsible for their glycosylation. In this study, a genome-wide analysis of the CitUGT gene family was conducted to identify CitUGTs that contribute to flavonoid 4'- O -glucosides biosynthesis. Our analysis identified 136 CitUGTs in the Citrus clementina genome, classifying them into 18 phylogenetic groups (A-R) and 25 families. This classification was strongly supported by consistent gene structures and motif patterns. Moreover, we identified a CitUGT gene ( Ciclev10025462m , designated CitUGT72AZ4 ) that encodes flavonoid 4'- O -glucosyltransferase for the first time in citrus. This enzyme preferentially glycosylated the 4'-OH group of multiple flavonoids, exhibiting higher catalytic efficiency toward quercetin and three flavones in vitro . Virus-induced gene silencing of CitUGT72AZ4 significantly decreased the accumulation of flavonoid 4'- O -glucosides. These results indicated that CitUGT72AZ4 participated in the biosynthesis of flavonoid 4'- O -glucoside in citrus. Overall, our findings provide valuable insights into the CitUGT gene family and contribute to its functional characterization.

The aim of this study was to explore the alterations in physicochemical characteristics, bioactivity, flavor and metabolic profiles of mango juice fermented by Limosilactobacillus reuteri FJG2526 (L. reuteri FJG2526). The results exhibited that L. reuteri FJG2526 had strong adaptability in mango juice, and reduced the total sugar, polyphenolics and flavonoids content of mango juice. L. reuteri FJG2526 fermentation ameliorated the flavor profiles of mango juice, particularly promoted the production of acids, alcohols, and esters. Moreover, 107 metabolites in the mango juice were drastically altered after 48 h L. reuteri FJG2526 fermentation by metabolomic analysis, including 73 remarkably upregulated metabolites and 34 remarkably downregulated metabolites, primarily involving amino acid metabolism. In addition, L. reuteri FJG2526 fermentation also enhanced the ability to scavenge DPPH and OH free radicals of mango juice, and inhibited lipase and α-glucosidase activities. This study offers new insights into the mango juice fermentation and will contribute to the application of L. reuteri in functional juices., 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 © 2025 Elsevier B.V. All rights reserved.)

Grape skin, consisting of a significant portion of grape pomace, is a great source of dietary fibers and polyphenols. In this study, grape skins of Hutai No. 8 were subjected to solid-state fermentation (SSF) using the fungal strain Eurotium cristatum. The total phenolic content (TPC) of grape skins increased and reached the maximum value after 8-day fermentation, which was 39.8% higher than those of the unfermented skins. The production of relevant enzymes was significantly enhanced, whereas the enzymatic activities of α-amylase and β-glucosidase showed significant positive correlations with the TPC. Sixteen phenolic compounds in the grape skins were identified and quantified using UPLC-QTOF-MS/MS and HPLC-DAD, whereas most of them increased significantly after fermentation. The antioxidant activities of grape skins showed a substantial improvement, which was positively and significantly correlated with TPC (p < 0.01). Principal component analysis indicated that the increased levels of ferulic acid, myricetin, resveratrol, and catechin during SSF were primarily responsible for the enhanced anti-1,1-diphenyl-2-picrylhydrazyl and anti-ABTS radical activities, as well as the ferric reducing ability of grape skins. Moreover, the changes in the concentrations of syringic acid, caffeic acid, gallic acid, protocatechuic acid, and quercetin-3-glucoside in grape skins exhibited a strong correlation with the ABTS radical scavenging capacity in vitro. Furthermore, fermentation time played a crucial role in the variation of phenolic profiles and antioxidant activities in grape skins. These findings indicate that SSF with E. cristatum effectively enhances the phenolic content in grape skin, thereby contributing to its antioxidant activities., (© 2025 Institute of Food Technologists.)

Cold stress adversely affects crop growth and productivity. Resolving the genetic basis of freezing tolerance is important for crop improvement. Wild potato (Solanum commersonii) exhibits excellent freezing tolerance. However, the genetic factors underlying its freezing tolerance remain poorly understood. Here, we identified flavonoid 3'-hydroxylase (F3'H), a key gene in the flavonoid biosynthesis pathway, as highly expressed in S. commersonii compared with cultivated potato (S. tuberosum L.). Loss of ScF3'H function impaired freezing tolerance in S. commersonii, while ScF3'H overexpression in cultivated potato enhanced its freezing tolerance. Metabolic analysis revealed that F3'H generates more downstream products by adding hydroxyl (-OH) groups to the flavonoid ring structures. These flavonoids enhance reactive oxygen species scavenging, thereby contributing to freezing tolerance. Furthermore, the W-box element in the F3'H promoter plays a critical role in cold responses. Cold-induced transcription factor ScWRKY41 directly binds to the ScF3'H promoter region and recruits histone acetyltransferase 1 (ScHAC1), which enhances histone acetylation at the F3'H locus and activates its transcription. Overall, we identified the cold-responsive WRKY41-F3'H module that enhances freezing tolerance by augmenting the antioxidant capacity of flavonoids. This study reveals a valuable natural gene module for breeding enhanced freezing tolerance in potato and other crops., Competing Interests: Conflict of interest statement. None declared., (© The Author(s) 2025. Published by Oxford University Press on behalf of American Society of Plant Biologists.)

Sorghum (Sorghum bicolor), one of the world's most important monocot crops, suffers severe yield losses due to attack by a polyphagous insect pest, fall armyworm (FAW; Spodoptera frugiperda). Here, we show that the Brown midrib 12 (Bmr12) gene, which encodes the caffeic acid O-methyltransferase (COMT) enzyme, promotes sorghum defense against FAW. Loss of Bmr12 function resulted in increased susceptibility, but enhanced resistance to FAW was observed in Bmr12-overexpression (OE) plants compared with wild-type (RTx430) plants. Although COMT is associated with modulating lignin levels, FAW infestation resulted in comparable lignin levels between bmr12 and Bmr12-OE sorghum plants. On the contrary, evidence presented here indicates that FAW feeding induced the accumulation of flavonoids, which was previously shown to have a negative impact on FAW growth and survival in Bmr12-OE plants compared with bmr12 and RTx430 plants. Furthermore, a combination of phytohormone profiling and transcriptomic analysis uncovered that COMT-mediated resistance to FAW depends on jasmonic acid (JA) and oxidative stress-associated pathways. Exogenous application of FAW oral secretions stimulated flavonoid accumulation in Bmr12-OE plants compared with bmr12 and RTx430 plants, indicating that COMT has an essential function in perceiving FAW oral cues. Taken together, the critical role of COMT in sorghum defense against FAW hinges upon the interplay between JA and its derivatives and hydrogen peroxide, which potentially helps to mount a robust flavonoid-based host defense upon caterpillar attack., Competing Interests: Conflict of interest statement. None declared., (© 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.)

Moderate UV-B promotes plant growth, but excessive UV-B inhibits plant development. The induction mechanism of how CtWD40-6 responds to UV-B is still unclear in safflower. Our results showed that CtWD40-6 is expressed at the top of safflower leaves and is strongly induced by UV-B. To further understand the function of the CtWD40-6 gene, we overexpressed the CtWD40-6 gene in safflower or Arabidopsis. First, different transgenic materials were treated with UV-B, and we found that the survival rate of plants overexpressing CtWD40-6 was significantly higher than that of the WT type. In contrast, the survival rate of wd40-6 mutant plants was significantly decreased compared with WT type. Then DAB, NBT and Trypan Blue staining were performed on different transgenic plants before and after UV-B treatment and the results showed that the staining of mutant and WT was significantly higher than that of overexpressing CtWD40-6. By comparing the data before and after UV-B stress, we found that the flavonoid content, antioxidant enzyme activity, chlorophyll content and photosynthetic rate of transgenic plants overexpressing CtWD40-6 were higher than those of WT and mutants, thereby obtaining better UV-B tolerance. Finally, we used yeast two-hybrid and luciferase complementation experiments to prove that CtWD40-6 increases the content of safflower flavonoids by interacting with CtANS1/CtCHS1/Ct4CL1/CtFLS1, thereby enhancing the plant's UV-B tolerance. The above results provide a theoretical basis for preliminary analysis of how safflower responds to UV-B stress through the transcriptional regulation of CtWD40-6., Competing Interests: Declaration of competing interest The authors declare that they have no competing interests., (Copyright © 2025 Elsevier Masson SAS. All rights reserved.)

Flavonoids constitute a class of natural compounds with varied bioactivities. Nevertheless, the potential health benefits of flavonoids for humans are often compromised by their low water solubility and limited bioavailability. In this study, four derivatives, namely 2',5'-dihydroxyflavanone (2), 5'-dihydroxyflavone-2'-O-β-d-glucuronide (3), and two isomers of hydroxyflavanone-2'-O-β-d-glucuronide (4 and 5), were biosynthesized from substrate 2'-hydroxyflavanone (1) through the specific hydroxylation and glucuronidation using Streptomyces coeruleorubidus NRRL B-2569. Product 2 was identified as a known compound while products 3-5 were structurally characterized as new structures through extensive 1D and 2D NMR analysis. The water solubility of obtained products 3-5 were enhanced by 36-340 times compared to the substrate. Moreover, the antioxidant assay revealed that product 3 exhibited improved 2,2-diphenyl-1-picrylhydrazyl (DPPH) radical scavenging activity compared to the substrate, decreasing the logIC 50 from 10.77 ± 0.05 μM to 9.55 ± 0.05 μM. Compound 3 also displayed significantly higher anticancer activity than the substrate 2'-hydroxyflavanone against Glioblastoma 33 cancer stem cells (GSC33), decreasing the IC 50 from 25.05 μM to 7.07 μM. Thus, S. coeruleorubidus NRRL B-2569 stands out as an effective tool for modifying flavonoids, thereby enhancing their water solubility and bioactivities., (Copyright © 2024 The Society for Biotechnology, Japan. Published by Elsevier B.V. All rights reserved.)

Jianpihuazhuotiaozhi granules (JPHZTZ) are traditional Chinese medicine formula. An analytical method using ultra-high-performance liquid chromatography coupled with quadrupole-time-of-flight mass spectrometry was established to characterize the chemical constituents of JPHZTZ and determine its metabolic profile in rat urine and plasma after oral administration. A total of 220 compounds were identified in JPHZTZ extract, including 61 flavonoids, 16 alkaloids, 53 organic acids, 31 terpenoids, and 59 other compounds. Among these compounds, 11 were tentatively identified by comparing the retention times and mass spectral data with the corresponding reference standards and the literature; the other 209 components were tentatively identified based on their mass spectra alone. After the oral administration of JPHZTZ extract to rats, 13 prototypes and 54 metabolites were identified or tentatively characterized based on their retention time and mass spectra. The primary in vivo metabolic reactions that occurred after the administration of JPHZTZ extract included glucuronidation, sulfation, hydroxylation, and methylation. The 13 prototypes and 54 metabolites were identified in rat urine and plasma and were confirmed to be the potential active ingredients of JPHZTZ. Our findings provide a starting point for the further elucidation of the mechanism of action of JPHZTZ., (© 2025 John Wiley & Sons Ltd.)

This study investigates the effects of silver nanoparticles (Ag NPs), biogenic silver nanoparticles derived from Rhizopus spp. (R.Ag NPs), and Rhizopus (R) elicitors on the yield and bioactive compounds of turmeric (Curcuma longa) using foliar spray and rhizome dipping techniques. Elicitors applied at concentrations of 10, 50, and 100 ppm over 210 days significantly enhanced turmeric yield, phenolic, flavonoid, antioxidant, and curcuminoid content compared to control plants. Among the treatments, Rhizopus culture at 50 ppm applied via foliar spray resulted in a 3.5% increase in curcuminoid content, while rhizome dipping at the same concentration led to a 3.75% increase. These findings suggest that foliar spraying and rhizome dipping with Rhizopus elicitors can effectively enhance turmeric quality, offering potential for improving crop production and curcuminoid yield., Competing Interests: Declaration of competing interest The authors declare the following financial interests/personal relationships which may be considered as potential competing interests:, (Copyright © 2025 Elsevier Masson SAS. All rights reserved.)

Lacquer tree (Toxicodendron vernicifluum) is an important economic crop and is rich in flavonoids. ACC (1-aminocyclopropane-1-carboxylic acid) is the precursor to ethylene. ACC treatment can induce physiological and biochemical responses in plants. In this study, flavonoids in the leaves of Toxicodendron vernicifluum, treated with ACC at different time points, were identified and analyzed. A total of 83 flavonoids were identified, including 38 flavonoids and 45 flavonoid glycosides. Among these, 48 flavonoids were first reported in T. vernicifluum. The total flavonoid content significantly increased following ACC treatment, although the accumulation patterns of individual flavonoids varied. Flavonoids with similar structure or glycosylation/acetylation modifications exhibited a positive correlation in their content changes in T. vernicifluum leaves under ACC treatment. Transcriptome sequencing was conducted on control and 3-days ACC-treated leaves, revealing an upregulation in the expression of genes related to flavonoids biosynthesis, such as PAL, CCR, CHS, MYB and ERF encoding genes. We hypothesized that ACC regulated flavonoids biosynthesis by activating ERF and MYB transcription factors in the ethylene signaling pathway. This study provided evidence for the regulation of flavonoids biosynthesis in lacquer trees through ACC treatment., 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 © 2025 Elsevier Masson SAS. All rights reserved.)

Flavonoids are the major medicinally active ingredients that exert potential effects in Amomum tsao-ko. In total, 277 flavonoid metabolites were identified in fresh and dried fruits of three different accessions of A. tsao-ko (Amomum tsao-ko), which could be classified into eight classes with more metabolites classified as flavonol. Furthermore, 193 differential flavonoid metabolites (DFMs) were selected from the six samples (three fresh and dried fruits), the main DFMs identified in the fresh and dried fruits of A. tsao-ko were apigenin and its derivatives, and 11 main DFMs were identified. Upon analyzing the variations in flavonoid content between fresh and dried fruits, our findings indicate that the L accession demonstrate a superior accumulation of flavonoid metabolites in their dried fruits. Combination with transcriptome data, the synthesis of flavone and isoflavone metabolites in fresh fruits of A. tsao-ko may be affected by the changes in the expression of chalcone isomerase genes and regulated by the NAC and AP2 transcription factor. This study provides an important theoretical basis for the functional study of flavonoid synthesis-related genes and transcription factors in A. tsao-ko fruits and for the breeding of these fruits with high flavonoid content., 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 © 2025 Elsevier B.V. All rights reserved.)

This study explored the changes in nutrients, metabolites, and enzyme activity in Citrus reticulata peel powders (CRPP) under conventional or ultrasound-assisted solid-state fermentation (SSF) using Aspergillus niger CGMCC 3.6189. Compared to nonfermented CRPP (NF-CRPP), ultrasound-assisted fermented CRPP (UIS-CRPP) significantly increased total protein and carotenoid levels by 85.26 % and 179.68 %, respectively, surpassing conventionally-fermented CRPP (FO-CRPP). Among the 521 identified differential metabolites, organic acids, lipids, and flavonoids were predominant. Flavonoid accumulation was primarily driven by the flavone and flavonol biosynthesis pathway, with 90.47 % and 90.00 % of differential flavonoids upregulated in FO-CRPP and UIS-CRPP, respectively. SSF significantly increased phenylalanine, tyrosine, and methionine levels, and tyrosine ammonia-lyase and β-D-glucosidase activities, with higher levels in UIS-CRPP. These findings suggest that conventional and ultrasound-assisted fermentation enhances flavonoid levels in CRPP by modulating key enzyme activities in flavonoid biosynthesis and biotransformation. Our study offers a feasible approach for producing value-added products from citrus peel waste., 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 © 2024 Elsevier Ltd. All rights reserved.)

Microplastics/nanoplastics are a top global environmental concern and have stimulated surging research into plant-nanoplastic interactions. Previous studies have examined the responses of plants to nanoplastic stress at various levels. Plant-specialized (secondary) metabolites play crucial roles in plant responses to environmental stress, whereas their roles in response to nanoplastic stress remain unknown. Here, we systematically examined the physiological and biochemical responses of Ginkgo biloba, a species with robust metabolite-driven defenses, to polystyrene nanoplastics (PSNPs). PSNPs negatively affected seedling growth and induced phytotoxicity, oxidative stress, and nuclear damage. Notably, PSNPs caused significant flavonoid accumulation, which enhances plant tolerance and detoxification against PSNP stress. To determine whether this finding is universal in plants, we subjected Arabidopsis, poplar, and tomato to PSNP stress and verified the common response of enhanced flavonoids across these species. To further confirm the role of flavonoids, we employed genetic transformation and staining techniques, validating the importance of flavonoids in mitigating excessive oxidative stress induced by NPs. Matrix analysis of transgenic plants with enhanced flavonoids further demonstrated altered downstream pathways, allocating more energy towards resilience against nanoplastic stress. Collectively, our results reveal the flavonoid multifaceted roles in enhancing plant resilience to nanoplastic stress, providing new knowledge about plant responses to nanoplastic contamination., (© 2024 Wiley‐VCH GmbH.)

The plant Polygonum capitatum (P. capitatum) contains a variety of flavonoids that are distributed differently among different parts. Nevertheless, differentially expressed genes (DEGs) associated with this heterogeneous distribution have not been identified. In this study, combined with transcriptomic and metabonomic analysis, we identified significant DEGs related to variations in flavonoid composition among different parts of P. capitatum. Subsequently, transcriptomic and nontargeted metabolomic analyses revealed that flavonoids and phenolic acids in different parts of P. capitatum were significantly enriched in the phenylpropanoid biosynthesis, shikimic acid biosynthesis, and flavonoid biosynthesis pathways. The expression levels of genes encoding enzymes, including shikimate O-hydroxycinnamoyltransferase (HCT), chalcone synthase (CHS), flavonoid 3',5'-hydroxylase (CYP75A), flavones 3-hydroxylase (F3H), flavonol synthase (FLS), leucoanthocyanidin reductase (LAR), trans-cinnamate 4-monooxygenase (CYP73A), and shikimate kinase (SK), were found to be the lowest in the leaves of P. capitatum via quantitative PCR. Interestingly, these genes are involved in the biosynthesis of quality markers such as gallic acid, quercetin, and quercitrin in P. capitatum. Finally, the targeted metabolomic results reconfirmed that the gallic acid, quercetin, and quercitrin contents were the highest in the leaves of P. capitatum. This research provides a theoretical basis for further understanding the differential regulatory mechanism of flavonoid metabolism in different parts of P. capitatum, providing novel insights into the pharmacognostic basis of P. capitatum., (© 2025 The Author(s). The Plant Genome published by Wiley Periodicals LLC on behalf of Crop Science Society of America.)

Drought stress poses a significant threat to global agricultural production, including the cultivation of medicinal plants. Plant growth-promoting bacteria (PGPB) and the eco-friendly element silicon (Si) are known to alleviate the adverse effects of drought stress. This study examines how inoculation with Bacillus pumilus G5 or/and Si influences plant hormone signaling and flavonoid biosynthesis pathways in drought-stressed Glycyrrhiza uralensis Fisch. (G. uralensis), focusing on genetic and metabolic aspects. The results indicate that the combined application of G5 and Si (G5+Si) may regulate the crosstalk among jasmonate (JA), gibberellin (GA), and ethylene (ET) signaling pathways, thereby up-regulating key flavonoid biosynthesis genes, including phenylalanine ammonia-lyase (PAL), cinnamate 4-hydroxylase (C4H), 4-coumarate-CoA ligase (4CL), and chalcone synthase (CHS), leading to the accumulation of isoliquiritigenin, liquiritigenin, liquiritin, and licochalcone A, thereby enhancing the drought tolerance of G. uralensis seedlings. The findings provide new insights into the synergistic role of PGPB and Si in improving plant resilience to drought stress, offering theoretical reference for further studies on plant drought tolerance mechanisms., 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 © 2025 Elsevier Masson SAS. All rights reserved.)

Hairy roots mediated by Agrobacterium rhizogenes can be obtained from the composite plants (plants with hairy roots and untransformed aerial parts) by ex vitro method. Composite plants can produce higher amounts of secondary metabolites by merging hydroponic systems. This provides a stable condition for composite plants, in which various metabolites are produced in different parts. In the present study, composite plants of Glycyrrhiza glabra were produced under ex vitro conditions and transferred into a hydroponic culture medium to produce and extract secondary metabolites. The results showed a 4.8- and 1.8-fold enhancement in the expressions of SQS1 and bAS genes in the roots of the composite plants compared to the control group, respectively. The levels of glycyrrhizin (1.7) and glabridin (3.5) were higher in the leaves of the composite plants compared to the controls. Moreover, higher amounts of glabridin (1.7) were observed in the roots of the composite plants compared to the control group. Investigation of the oxidative enzymes in the composite plants and control group revealed that the plants used more secondary metabolites through Agrobacterium inoculation. The plants needed more antioxidant enzymes to counter the release of oxygen-free radicals in control conditions, but composite plants used secondary metabolites as scavengers. Data revealed that composite plants managed to produce high amounts of various secondary metabolites in a hydroponic system., Competing Interests: Declaration of Competing Interest The authors declare no financial or commercial conflicts of interest., (Copyright © 2025. Published by Elsevier B.V.)

Clostridium butyricum inhabits various anoxic environments, including soil and the human gut. Here, this common bacterium comes into contact with abundant plant-derived flavonoids. Metabolization of these bioactive polyphenols has been studied in recent years, particularly focusing on gut bacteria due to the proposed health-promoting properties of these dietary constituents. Based on an initial report in 1997 on eriodictyol degradation (Miyake et al. 1997, J Agric Food Chem, 45:3738-3742), the present study systematically investigated C. butyricum for its ability to convert a set of structurally diverse flavonoids. Incubation experiments revealed that C. butyricum deglycosylated flavonoid O-glucosides but only when glucose was absent. Moreover, aglycone members of flavone, flavanone, dihydrochalcone, and flavanonol subclasses were degraded. The C-ring cleavage of the flavanones, naringenin and eriodictyol, was stereospecific and finally resulted in formation of the corresponding hydroxyphenylpropionic acids. Stereospecific C-ring cleavage of the flavanonol taxifolin led to taxifolin dihydrochalcone. C. butyricum did neither cleave flavonols and isoflavones nor catalyze de-rhamnosylation, demethylation, or dehydroxylation of flavonoids. Genes encoding potential flavonoid-metabolizing enzymes were detected in the C. butyricum genome. Overall, these findings indicate that C. butyricum utilizes flavonoids as alternative substrates and, as observed for the dihydrochalcone phloretin, can eliminate growth-inhibiting flavonoids through degradation. KEY POINTS: • Clostridium butyricum deglycosylated flavonoid O-glucosides. • Clostridium butyricum converted members of several flavonoid subclasses. • Potential flavonoid-metabolizing enzymes are encoded in the C. butyricum genome., Competing Interests: Declarations. Ethics approval: This article does not contain any studies involving human participants or animals performed by the author. Conflict of interest: The author declares no competing interests., (© 2025. The Author(s).)

Light is a crucial factor in plant growth and development. Plants exposed to light stress experience various effects on their growth. This research was conducted to investigate the effects of different light intensities on morpho-physiological traits, phytochemical compounds, and gene expression related to the biosynthesis of voletile in Thymus vulgaris L. The results demonstrated that light intensity (20, 50, 70 and 100%) had a significant impact on morpho-physiological characteristics, pigments content, antioxidant enzymes activities, as well as the content of MDA, H2O2, anthocyanin, thymol, carvacrol, phenols, flavonoids, essential oils, and monoterpenes. Moreover, the expression of the biosynthesis genes of monoterpene compounds was significantly influenced by light intensity. While an increase in light intensity led to higher leaf count (164.6%) and biomass (33.5%), it was accompanied by a decrease in leaf area, stem length, and internode length. The highest levels of chlorophyll a (4.92 mgg-1 FW) and b (1.75 mgg-1 FW), carotenoids (907.31 µ Mg-1FW), MDA (9.93 µ Mg-1FW), anthocyanin, SOD (29.62 Umg - 1 Protein), thymol (41.2%), and carvacrol (4.46%) were observed at 70% treatment and decreased as light intensity increased. Also, H2O2, catalase and polyphenol oxidase activities, phenols, flavonoids, essential oils, and monoterpenes increased with higher light intensity, with the highest H2O2 concentration recorded at 100% (4.43 fold). Importantly, key genes involved in monoterpene biosynthesis, including DXR, TPS, CYP71D178, and CYP71D179, exhibited significantly enhanced expression under full light conditions compared to other light intensities. In conclusion, increased light intensity stimulated the elevation of oxidative indicators, antioxidant activity and enhancing the expression of genes involved in phytochemical compound biosynthesis and consequently leading to the accumulation of volatile compounds in Thymus vulgaris L. Future research will focus on investigating the combined effects of various abiotic stresses at the field level and extending the stress duration to evaluate potential additive effects., Competing Interests: NO authors have competing interests., (Copyright: © 2025 Hashemifar et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

The phenylpropanoid pathway, regulated by transcription factors of the MYB family, produces secondary metabolites that play important roles in fertilization and early phase of fruit development. The MYB46 transcription factor is a key regulator of secondary cell wall structure, lignin and flavonoid biosynthesis in many plants, but little is known about its activity in flowers and berries in F. vesca. For functional analysis of FvMYB46, we designed a CRISPR-Cas9 construct with an endogenous F. vesca-specific U6 promoter for efficient and specific expression of two gRNAs targeting the first exon of FvMYB46. This generated mutants with an in-frame 81-bp deletion of the first conserved MYB domain or an out-of-frame 82-bp deletion potentially knocking out gene function. In both types of mutant plants, pollen germination and fruit set were significantly reduced compared to wild type. Transcriptomic analysis of flowers revealed that FvMYB46 positively regulates the expression of genes involved in processes like xylan biosynthesis and metabolism, homeostasis of reactive oxygen species (ROS) and the phenylpropanoid pathway, including secondary cell wall biosynthesis and flavonoid biosynthesis. Genes regulating carbohydrate metabolism and signalling were also deregulated, suggesting that FvMYB46 might regulate the crosstalk between carbohydrate metabolism and phenylpropanoid biosynthesis. In the FvMYB46-mutant flowers, the flavanol and flavan-3-ol contents, especially epicatechin, quercetin-glucoside and kaempferol-3-coumaroylhexoside, were reduced, and we observed a local reduction in the lignin content in the anthers. Together, these results suggest that FvMYB46 controls fertility and efficient fruit set by regulating the cell wall structure, flavonoid biosynthesis, carbohydrate metabolism, and sugar and ROS signalling in flowers and early fruit development in F. vesca., Competing Interests: Declarations. Ethics approval and consent to participate: Not applicable. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Flavonoids are key bioactive components for evaluating the pharmacological activities of Chimonanthus praecox . Exploring the potential flavonoids and pharmacological mechanisms of C . praecox lays a foundation for the rational development and efficient utilization of this plant. This study employed ultra-performance liquid chromatography-tandem mass spectrometry-based widely targeted metabolomics to comprehensively identify the flavonoids in C . praecox . Network pharmacology was employed to explore the bioactive flavonoids and their mechanisms of action. Molecular docking was adopted to validate the predicted results. Finally, the content of bioactive flavonoids in different varieties of C . praecox was measured. The widely targeted metabolomics analysis identified 387 flavonoids in C . praecox , and the flavonoids varied among different varieties. Network pharmacology predicted 96 chemical components including 19 bioactive compounds, 181 corresponding targets and 2 504 disease targets, among which 99 targets were shared by the active components and the disease. Thirty-three core targets were predicted, involving 229 gene ontology terms and 99 pathways ( P ≤0.05), which indicated that the flavonoids components of C . praecox exhibited pharmacological activities including antioxidant, anti-inflammatory, antimicrobial, and antiviral activities. Topological analysis screened out five core components (salvigenin, laricitrin, isorhamnetin, quercetin, and 6-hydroxyluteolin) and five core targets (SRC, PIK3R1, AKT1, ESR1, and AKR1C3). The predicted bioactive flavonoids from C . praecox stably bound to key targets, which indicated that these flavonoids possessed potential bioactivities in their interactions with the targets. The flavonoids in C . praecox exerted pharmacological activities in a multi-component, multi-target, and multi-pathway manner. The combined application of metabolomics and network pharmacology provides a theoretical basis for in-depth studies on the pharmacological effects and mechanisms of C . praecox .

Ophiopogon japonicus , a precious medicinal plant endemic to Zhejiang Province. Its tuberous roots are rich in bioactive components such as flavonoids, possessing anti-inflammatory, antioxidant, and immunomodulatory properties. To elucidate the impact of cadmium (Cd) stress on the accumulation and biosynthetic pathway of flavonoids in O . japonicus , this study exposed O . japonicus to different concentrations of Cd stress and explored the changes through integrated transcriptomics and metabolomics analysis. The results demonstrated that Cd stress (1 mg/L and 10 mg/L) significantly increased the content of flavonoids in O . japonicus in a concentration-dependent manner. The metabolomics analysis revealed a total of 110 flavonoids including flavones, flavanols, flavonols, flavone and flavonol derivatives, flavanones, isoflavonoids, chalcones and dihydrochalcones, and anthocyanins in O . japonicus , among which flavones, flavonols, flavone and flavonol derivatives, and anthocyanins increased under Cd stress. The transcriptomics analysis identified several key flavonoid biosynthesis-associated genes with up-regulated expression under Cd stress, including 14 genes encoding 4-coumarate CoA ligase (4CL), 2 genes encoding chalcone isomerase (CHI), and 14 genes encoding phenylalanine ammonia lyase (PAL). The gene-metabolite regulatory network indicated significant positive correlations of 4CL ( Cluster-21637 . 5012 , Cluster-21637 . 90648 , and Cluster-21637 . 62637 ) and CHI ( Cluster-21637 . 111909 and Cluster-21637 . 123300 ) with flavonoid metabolites, suggesting that these genes promoted the synthesis of specific flavonoid metabolites, which led to the accumulation of total flavonoids under Cd stress. These findings provide theoretical support for the cultivation and utilization of medicinal plants in Cd-contaminated environments and offered new perspectives for studying plant responses to heavy metal stress.

Key Message: Blue light influences the MYB gene family, resulting in varying accumulations of different flavonoids in rice caryopsis at distinct developmental stages, with a higher concentration observed in the initial stage. The regulatory effect of blue light on plant flavonoids has been extensively documented; however, its influence on the development of rice caryopsis morphology remains unreported. Through the analysis of transcriptomes, proteomes, and metabolites, combined with Weighted Gene Co-expression Network Analysis (WGCNA), the accumulation of flavonoids in rice caryopsis under blue light at various developmental stages was thoroughly examined. Furthermore, four MYB family transcription factors (TFs) that significantly influence the structural genes involved in flavonoid biosynthesis were identified. The results indicate that the accumulation of flavonoids primarily occurs during the early stages of caryopsis development. Key structural genes, including PAL, 4CL, CHS, CHI, F3H, and FLS, are upregulated in both gene and protein expression when exposed to blue light. Moreover, the WGCNA analysis identified several TFs that may influence these genes, including Os08t0144000-01 and Os01t0695900-01, as well as the proteins Q7F3D6, Q2QM89, A0A0P0W9C3, and Q6ZDM0, all of which belong to the MYB family. The research has enhanced our understanding of flavonoid accumulation in rice caryopsis when exposed to blue light. It also establishes a framework for the synthesis of secondary metabolites induced by blue light, thereby creating more opportunities to enhance the quality of horticultural plants., Competing Interests: Declarations. Conflict of interest: The authors declare that they have no competing interests. Ethical approval and consent to participate: Not applicable. Consent for publication: Not applicable., (© 2025. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Rosehip (Rosa canina L.) is a perennial medicinal plant from the Rosaceae family. Due to its important bioactive compounds and oil, its use in the food, pharmaceutical, and cosmetic industries is expanding. As elicitation is an effective strategy for the production of secondary metabolites and oil, this study aimed to investigate the exogenous application of salicylic acid (SA) (0, 1, 2, and 3 mM) and jasmonic acid (JA) (0, 0.5, 1, and 1.5 mM) on the accumulation and production of oil, the profile of seed fatty acids, as well as the content of phenolic acids, vitamin C, total carotenoid, anthocyanin, total phenolic content, and total flavonoid content of rosehip pericarp. Based on the results, applying SA and JA externally increased the seed oil content by 32.57% compared to the control. The treatment of 3 mM SA and 1.5 mM JA resulted in the highest (11.68%) seed oil content. Eicosanoic acid and palmitic acid production increased under this treatment, while the amount of linoleic acid decreased as the concentrations of SA and JA increased. There was a significant increase in the amount of phenolic acids in rosehip pericarp extract under the influence of SA and JA treatments. Spraying with 3 mM SA and 1.5 mM JA increased vitamin C content and total phenol content by 50.44% and 39.13%, respectively, compared to the control. Additionally, the treatment of 2 mM SA and 1.5 mM JA resulted in the highest total flavonoid content, antioxidant activity, and total carotenoid. These results suggest that using appropriate concentrations of SA and JA as biodegradable, fast, and cost-effective stimulants can be a suitable solution for increasing the production of seed oil content and secondary metabolites of rosehip extract on a large scale, supplying raw material for pharmaceutical, cosmetic, and food industries., Competing Interests: Declarations. Ethics approval and consent to participate: This manuscript is an original research and has not been published or submitted in other journals. Ethical review: This study does not involve any human or animal testing. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Background: Chalcone synthase (CHS) is a key rate-limiting enzyme in the flavonoid synthesis pathway. Flavonoids are crucial secondary metabolites that play significant roles in plant growth, development, and stress resistance. The CHS gene (BrCHS) family in Chinese cabbage has not yet been studied., Results: We identified 10 BrCHS genes distributed across 7 chromosomes in the Chinese cabbage genome. Their encoded proteins all contain the Chal&#95;Sti&#95;Synt&#95;C (PF02797) and Chal&#95;Sti&#95;Synt&#95;N (PF00195) domains and can be classified into two groups based on systematic evolution analysis. These BrCHS genes contain 2-4 exons and numerous cis-acting elements responsive to light, hormones, stress, growth and development in the BrCHS gene promoters. We also revealed that the expression of BrCHS2 and BrCHS8 increased under treatment with methyl jasmonate, salt, or drought stress. Virus-induced gene silencing (VIGS) of BrCHS4 inhibited the expression of BrCHS4 and reduced the flavonoid and anthocyanin contents in leaves., Conclusions: Ten BrCHS family genes are present in the genome of Chinese cabbage. These BrCHS genes seemingly maintained similar characteristics and functionalities during evolution. Our results demonstrated that BrCHS4 is involved in flavonoid and anthocyanin accumulation in Chinese cabbage and identified candidate genes for purple Chinese cabbage breeding., Competing Interests: Declarations. Ethics approval and consent to participate: Not applicable. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Cupressus torulosa , an evergreen tree commonly known as the Himalayan or Bhutan cypress, is a significant coniferous species native to the Himalayan regions of Bhutan, northern India, Nepal, and Tibet. In this study, we employed ultra-performance liquid chromatography quadrupole time-of-flight mass spectrometry (UPLC-QTOF-MS) integrated with chemometrics to analyze the metabolite profiles of C. torulosa needles collected from 14 distinct geographical regions. Advanced statistical tools, including Principal Component Analysis (PCA) and Hierarchical Cluster Analysis (HCA), were utilized to identify significant variations in the chemical composition across these locations. Our analysis identified 24 marker compounds consistently present in all samples, screened using stringent filtering criteria including One-way ANOVA, moderated T -test, and multiple testing correction with the Benjamini-Hochberg FDR method. Additionally, for the first time, we established the flavonoid biosynthesis pathway in C. torulosa , enhancing the understanding of its metabolic fingerprint. These findings provide critical insights into the phytochemical diversity of C. torulosa and offer valuable tools for quality control, authentication and advancing the application of UPLC-QTOF-MS in natural product research.

Phenylalanine (Phe), an aromatic amino acid, is a key precursor of flavonoids, which are crucial for plant growth and development. Arogenate dehydratase (ADT) catalyzes the final step in Phe biosynthesis. This study identified eleven ADT genes in G. hirsutum, twelve in G. barbadense, six in G. arboreum, and six in G. raimondii. Among them, GhADT5 exhibited the highest upregulation under alkali stress. Silencing GhADT5 using virus-induced gene silencing (VIGS) reduced cotton tolerance to alkali stress. GhADT5 silencing also led to decreased plant phenylalanine content, total flavonoid content, and activities of superoxide dismutase (SOD), catalase (CAT), and peroxidase (POD). These reductions caused intracellular accumulation of Malondialdehyde (MDA) and reactive oxygen species (ROS). This oxidative damage ultimately reduced tolerance to alkali stress. In addition, silenced plants displayed reduced stomatal aperture, cellular deformation, and irregular intercellular breaks in the leaf epidermis. In summary, these findings suggest that GhADT5 may enhance resistance to alkali stress by regulating enzymatic and non-enzymatic antioxidant systems. This study highlights the role of GhADT5 under alkali stress and provides novel insights for breeding cotton varieties with improved stress tolerance., Competing Interests: Declarations. Ethical approval and consent to participate: All the cotton materials were collected from the Institute of Cotton Research, Chinese Academy of Agricultural Sciences, which are publicly and available for non-commercial purpose. All experimental studies on plants were complied with relevant institutional, national, and international guidelines and legislation. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Black and red rice have many physiological effects, including antioxidant capacity and decreasing blood glucose, because they are flavonoid-rich. Here, our study found that Hongcaomi (a red rice variety) of five examined rice cultivars contained the highest total polyphenol content and total flavonoid content and the strongest antioxidant activity, with values of 5.83 mg GAE/g, 1.57 mg RE/g, and 72.63 μmol Trolox/g, respectively. The decreased blood glucose effect showed that the strongest effect was shown by the two red rice cultivars, followed by the black rice cultivar Ashamurasaki, the white rice cultivar Yixiang1B, and the black rice cultivar Chongxuehei which had no effect. The study further found that the strongest physiological effect in red rice was derived from catechin. In addition, our study reveals that pigmented rice could decrease blood glucose levels through many pathways, including repairing liver morphology, improving the expression level of genes involved in gluconeogenesis and glycogenesis pathways, and regulating the composition of the gut microbiota.

Wintersweet ( Chimonanthus praecox ) is known for its flowering in winter and its rich floral aroma; the whole flower is yellow and the inner petals are red. In this study, we chose the wintersweet genotypes HLT040 and HLT015 as the research materials, and studied the co-regulatory mechanism of color and fragrance of wintersweet through metabolomics and transcriptomics. This study found that there were more flavonoids in HLT015, and anthocyanins (cyanidin-3-O-rutinoside and cyanidin-3-O-glucoside) were only present in HLT015, but HLT040 contained more monoterpenes and FVBPs (phenylpropanoid volatile compounds) than HLT015. We constructed putative benzenoids and phenylpropanoid metabolism pathway as well as terpene metabolism pathways. We found some linkages between the different structural genes and metabolites for flower color and fragrance in wintersweet, and screened out 39 TFs that may be related to one or more structural genes in benzenoids and phenylpropanoid or terpene metabolism pathways. In the yeast one-hybrid assay, we found that CpERF7 was able to interact with the promoter of CpANS1 , while CpbHLH50 and CpMYB21 interacted with the promoter of CpTPS4 . This study provides a theoretical basis for understanding the co-regulatory mechanism of color and fragrance in wintersweet.

The effect of endophytic fungal biotransformation on the formation of metabolic substances has become a non-negligible factor in assessing the quality of medicinal plants. However, the relevant evidences are still particularly lacking. In this study, an endophytic fungus CA3-A with biotransformation activity was screened and identified as Talaromyces coprophilus from root of Astragalus mongholicus. Its biotransformation effect on host metabolism was studied by co-culture with fungus and host root. The results showed that in addition to the significant changes in primary metabolites, secondary metabolite conversions are mainly focused on compounds such as terpenoids, phenols and flavonoids. The transformed metabolic pathways are mainly enriched in glycerophospholipid metabolism, ubiquinone and terpenoid quinones biosynthesis, and tyrosine metabolism. A total of 38 signature differential metabolites are found through multivariate statistical analysis. In addition to eleven primary metabolites including glycerides and glycerophospholipids, twenty-eight secondary metabolites distribute in terpenes, sterols, phenols, flavonoids, benzene, lipids and other classes of compounds, respectively. In conclusion, this study aims to quickly focus on the signature differentially metabolites from a large amount of information, narrow the possible range of potential transformation products, and provide help for understanding of influence of endophytes on host metabolism and the search for new natural products., Competing Interests: Declarations. Ethical Approval: not applicable. Consent to Participate: This article has been approved by all the authors. Consent to Publish: not applicable. Competing Interests: The authors declare no competing interests., (© 2025. The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature.)

Cadmium (Cd) is easily absorbed by rice and enters the food chain, posing a health risk to humans. Plant growth promoting bacteria (PGPB) can help the plant respond to Cd stress, but the mechanism of PGPB for Cd reduction is unclear. Therefore, this study was conducted and found inoculation with a newly isolated Pseudomonas koreensis promoted the growth of rice and reduced its Cd content. Fluorescent staining using PI and H 2 O 2 probe indicated that PGPB attenuated oxidative damage in rice. Metabolomics revealed that 59 metabolites were upregulated after inoculation, with phenylpropanoids and flavonoids being significantly activated. Spectrophotometry analysis comfirmed the content of flavonoid, lignin, phenol, glutathione, proline and the activities of antioxidant enzymes were higher in the inoculated rice than in the control. Quantitative PCR showed the expression of genes related to phenylpropanoids (OsPAL, OsC4H, Os4CL) and flavonoids (OsCHS, OsCHI) was significantly increased by PGPB, while the genes of heavy metal transporters (OsNRAMP5, OsHMA2, OsIRT1) were significantly decreased. Overall, this study provides an insight into the PGPB-mediated detoxification mechanism in rice under Cd stress and emphasizes the role of phenylpropanoids and flavonoids in the production of low-Cd rice to ensure human health., 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 © 2024 Elsevier B.V. All rights reserved.)

Pine wilt disease (PWD), caused by the pine wood nematode (PWN) Bursaphelenchus xylophilus, threatens Pinus seriously. Pinus koraiensis is one of the most important pine species in China and is the host for PWN. However, our understanding of the defence-regulating process following infection by B. xylophilus at the molecular level remains limited. To understand the mechanisms that P. koraiensis responds to B. xylophilus invasion, P. koraiensis was inoculated with B. xylophilus solutions and observed no obvious symptoms during the early stage; symptoms began to appear at 5 dpi. Therefore, we conducted comparative transcriptomic, metabonomic and proteomic analyses between P. koraiensis 5dpi and 0 dpi. In infected plants, 1574 genes were significantly up-regulated, including 17 terpenoid-, 41 phenylpropanoid- and 22 flavonoid-related genes. According to GO and KEGG enrichment analyses of significantly up-regulated genes, 86 GO terms and 16 KEGG pathways were significantly enriched. Most terms and pathways were associated with terpenoid-, phenylpropanoid-, flavonoid- and carbohydrate-related events. Similarly, the abundance of 36 and 30 metabolites, significantly increased in positive and negative polarity modes, respectively. Among them, naringenin and 3-methyl-2-oxovaleric acid exhibited significant toxic effects on B. xylophilus. According to functional analysis of significantly up-regulated metabolites, most terms were enriched in above pathways, in addition to alkaloid biosynthesis. Although the abundance of few proteins changed, response to stress term was significantly enriched in significant up-regulated proteins. Furthermore, plant receptor-like serine/threonine kinases, pectin methylation modulators, pinosylvin O-methyltransferase and arabinogalactan/proline-rich proteins were significantly up-regulated in the infected P. koraiensis compared to healthy plants. These proteins were not abundant in the healthy plant. Overall, these results indicate that P. koraiensis can actively response to PWN via various defense strategies, including events related to terpenoids, flavonoids, phenylpropanoids, lipids and alkaloids. Particularly, terpenoids and flavonoids are required for the early defence of P. koraiensis against B. xylophilus infection., Competing Interests: Declarations. Ethics approval and consent to participate: No specific permits were needed, and material collection and molecular experiments were performed in accordance with current Chinese regulations. Consent for publication: Not applicable. All authors agreed to publish. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Proanthocyanidins (PAs) play significant roles in plants and are bioactive compounds with health benefits. The polymerization mechanism has been debated for decades. Here we show that laccases (LACs) are involved in PA polymerization and miR397a is a negative regulator of PA biosynthesis in Salvia miltiorrhiza and Populus trichocarpa. Elevation of miR397a level causes significant downregulation of LACs, severe reduction of polymerized PAs, and significant increase of flavan-3-ol monomers in transgenic S. miltiorrhiza and P. trichocarpa plants. Enzyme activity analysis shows that miR397a-regulated SmLAC1 catalyzes the polymerization of flavan-3-ols and the conversion of B-type PAs to A-type. Both catechin and epicatechin can serve as the starter unit and the extension unit during PA polymerization. Overexpression of SmLAC1 results in significant increase of PA accumulation, accompanied by the decrease of catechin and epicatechin contents. Consistently, CRISPR/Cas9-mediated SmLAC1 knockout shows the opposite results. Based on these results, a scheme for LAC-catalyzed PA polymerization is proposed. The work provides insights into PA polymerization mechanism., Competing Interests: Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Juglans sigillata Dode is rich in flavonoids, but the low ratio of female to male flower buds limits the development of the J. sigillata industry. While the abundance of flavonoids in J. sigillata is known, whether flavonoids influence female flower bud differentiation has not been reported. In this study, we explored the regulatory mechanisms of gene expression and metabolite accumulation during female flower bud differentiation through integrated transcriptomic and metabolomic analyses. Our findings revealed that flavonoid biosynthesis is a key pathway influencing female flower bud differentiation, with metabolites primarily shifting towards the isoflavonoid, flavone, and flavonol branches. Structural genes such as chalcone synthase, dihydroflavonol 4-reductase, flavonol synthase, and flavonoid 3',5'-hydroxylase were identified as playing crucial regulatory roles. The expression of these genes promoted the accumulation of flavonoids, which in turn influenced female flower bud differentiation by modulating key regulatory genes including Suppressor of Overexpression of Constans1, Constans, Flowering Locus T, and APETALA1. Furthermore, transcription factors (TFs) highly expressed during the physiological differentiation of female flower buds, particularly M-type MADS, WRKY, and MYB, were positively correlated with flavonoid biosynthesis genes, indicating their significant role in the regulation of flavonoid production. These results offer valuable insights into the mechanisms of female flower bud differentiation in J. sigillata and highlight the regulatory role of flavonoids in plant bud differentiation., Competing Interests: Declarations. Ethics approval and consent to participate: All local, national or international guidelines and legislation were adhered to in the production of this study. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Rhaponticum carthamoides (Willd.) Iljin. is an endemic plant species found in Siberia, Mongolia, and Kazakhstan. Its roots and rhizomes are used to treat physical fatigue and weakness following illness. The present study examines the scaling up of caffeoylquinic acid (CQA) derivative and flavonoid production in R. carthamoides transformed roots. The transformed roots were grown in shaken Erlenmeyer flasks of varying volumes (0.5-2 L), a temporary immersion system (TIS) (Rita ® and PlantForm bioreactors), and a nutrient sprinkle bioreactor (NSB) in Woody Plant medium for 35 days. The highest dry biomass production was achieved in the 0.5 L and 1 L flasks and in the NSB bioreactor, yielding 22.2 to 20.4 g/L-approximately 14 to 23 times the weight of the inoculum. The accumulation of individual specialized metabolites varied depending on the culture system used. The peak amount of CQAs (544.5 mg/L), in terms of the increase in dry weight and metabolite levels, was obtained in the NSB bioreactor. The primary CQAs were chlorogenic acid (5-CQA) and a tri-CQA 1. The highest concentration of 5-CQA (7.38 mg/g DW) was found in the roots cultivated in the NSB bioreactor. In contrast, the tri-CQA 1 dominated in the roots from 2 L shaken Erlenmeyer flasks (8.44 mg/g DW). Our findings demonstrate that transformed roots growing in an NSB bioreactor are an effective system for increasing CQA production, potentially serving as an alternative source. This biotechnological approach could help reduce the overexploitation of field-grown R. carthamoides , a currently threatened species.

Plant growth behavior is a function of genetic network architecture. The importance of root microbiome variation driving plant functional traits is increasingly recognized, but the genetic mechanisms governing this variation are less studied. Here, we collect roots and rhizosphere soils from nine Populus species belonging to four sections (Leuce, Aigeiros, Tacamahaca, and Turanga), generate metabolite and transcription data for roots and microbiota data for rhizospheres, and conduct comprehensive multi-omics analyses. We demonstrate that the roots of vigorous Leuce poplar enrich more Pseudomonas, compared with the poorly performing poplar. Moreover, we confirm that Pseudomonas is strongly associated with tricin and apigenin biosynthesis and identify that gene GLABRA3 (GL3) is critical for tricin secretion. The elevated tricin secretion via constitutive transcription of PopGL3 and Chalcone synthase (PopCHS4) can drive Pseudomonas colonization in the rhizosphere and further enhance poplar growth, nitrogen acquisition, and secondary root development in nitrogen-poor soil. This study reveals that plant-metabolite-microbe regulation patterns contribute to the poplar fitness and thoroughly decodes the key regulatory mechanisms of tricin, and provides insights into the interactions of the plant's key metabolites with its transcriptome and rhizosphere microbes., Competing Interests: Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Background: Camellia oleifera Abel. is one of the four major woody oil species whose seeds produce high-grade edible oil. In recent years, the planting area of Camellia oleifera is increasing in China. However, in the process of cultivation, due to the high fruit load, the Camellia oleifera tree has small fruit and poor quality. In previous studies, we explored the optimal leaf-fruit ratio. In this study, the changes of molecular regulation mechanism of Camellia oleifera under different leaf-fruit ratios were revealed by combining physiological indexes with transcriptome data., Result: The physiological results showed that the content of MDA and starch in leaves increased significantly with the decrease in the leaf-to-fruit ratio. The results of transcriptome showed that there was a close relationship between leaf-fruit ratio and phenylpropanoid biosynthesis pathway. With the decrease of leaf-fruit ratio, the expression of genes related to lignin and flavonoid biosynthesis increased significantly, which promoted the synthesis of lignin and flavonoid., Conclusions: Combining physiological indicators and transcriptomics, we demonstrated that leaf-fruit ratio can significantly affect the normal growth of plants. When the fruit load is too high, the fruit as a 'sink' will consume a large amount of nutrients in the plant body and promote the biosynthesis of lignin and flavonoids in the body. The results provide a more reliable scientific basis for the study of cultivation and management of Camellia oleifera., Competing Interests: Declarations. Ethics approval and consent to participate: The plant samples used in this study were grown and collected at Central South University of Forestry and Technology in Hunan, China. The samples were collected in strict accordance with the relevant regulations. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Postharvest pericarp browning, caused primarily by the enzymatic oxidation of phenols, reduces the shelf life and market value of litchi fruit and is considered a major limitation for the development of the litchi industry. Previous studies have shown that polyphenol oxidase (PPO) is a key enzyme and that flavonoids are important substrates for enzymatic browning; however, direct evidence is still lacking. This study investigated the differences in the browning process among the wild type (WT) and four PPO gene-edited litchi calli to verify the function of PPO in the browning of litchi tissues. Compared to the WT callus, the proliferation rate, relative expression of litchi PPO gene (LcPPO), PPO activity and color changes significantly decreased or slowed down in all gene-edited calli, indicating that the latter exhibited a slower browning process. Using a liquid chromatography tandem mass spectrometry approach (LC-MS/MS), 83 metabolites of flavonoids were identified, of which 58 were differentially accumulated metabolites (DAMs). Venn analysis revealed 12 common DAMs across different genotypic contrasts that were mostly enriched in the flavonoid biosynthesis pathway. It was presumed that the decrease of LcPPO expression in gene-edited calli led to the reduced PPO activity, then reduced the (-)-epicatechin oxidation. The accumulation of (-)-epicatechin caused the common upregulation of procyanidin B2 and upstream substances such as dihydrokaempferol, taxifolin, naringenin chalcone, 7,4'-dihydroxyflavone, and rutin in their biosynthesis pathways. The results provide novel evidence that (-)-epicatechin acts as the primary direct substrate in the enzymatic browning reaction mediated by PPO., 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 © 2024. Published by Elsevier B.V.)

The impact of drought stress on plant growth, development, and productivity presents a significant challenge in various environments worldwide. The exogenous application of polyamines as osmotically active materials plays a crucial role in enhancing plant tolerance to environmental stress. In this study, we examined the effects of L- and D-enantiomers of ornithine (0 and 1 mM) under both well-watered and drought stress conditions on the growth traits, essential oil (EO) yield, and composition, gene expression, and total phenolic and flavonoid content of Salvia officinalis. The experiment was designed as a factorial experiment using a completely randomized design with three replications. The results demonstrated that drought stress led to a decrease in plant biomass and an increase in EO content, chemical profiles of the EO, and total phenolic and flavonoid content compared to the respective control values. However, the exogenous supplementation of ornithine particularly D-ornithine resulted in enhanced stem, leaf, and total plant biomass, a 20% increase in EO content, and a 75% increase in yield. Additionally, these were increases of 11.76% in total phenol and 70%, 105.66%, and 114.28% in flavonoid content when compared to well-watered plants without ornithine supplementation. These improvements were strongly linked to growth enhancement, as evidenced by principal component analysis (PCA). The EO extracted from S. officinalis consisted of 22 compounds, primarily monoterpenes, including α-thujone (18.47-41.65%), camphor (15.05-25.17%), 1,8-cineole (10.12-21.6%), and β-thujone (6.23-21.2%). The percentage of these volatile compounds was found to be highest in D-ornithine-treated stressed plants compared to control conditions. The interaction between water availability and the application of D-ornithine and L-ornithine significantly influenced the expression of borneol diphosphate synthase (BS), sabinene synthase (SS), and cineole synthase (CS) under drought stress, with notable upregulation observed compared to normal growth conditions. Specifically, D-ornithine enhanced the expression of BS and SS by 45.29% and 113.63%, respectively, under drought stress, while both D-ornithine and DL-ornithine significantly increased CS expression. The present results suggest that D-ornithine may serve as a stress-protecting compound, increasing total phenol and flavonoids content, thereby enhancing the capacity of the antioxidant system and increasing EO compounds under drought stress., Competing Interests: Declarations. Ethics approval and consent to participate: The one-year-old mother plants of Salvia officinalis used in this study were collected with the approval of local regulatory authorities. All methods were carried out in compliance with regional and national regulations. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Luohan Guo (Siraitia grosvenorii) is a traditional food homologous fruit in China, which is famous owing to its rich natural antioxidant secondary metabolites. However, the impacts of ripening stages and edible parts on high-resolution metabolic profiles of Luohan Guo are poorly understood till now, greatly limiting its comprehensive utilization. In this study, an LC-QTOF/MS-based non-targeted metabolomics approach was carried out to reveal the dynamic accumulation of metabolites in the peel and pulp of Luohan Guo across seven growth stages, and further revelation their potential pharmacological activities and mechanisms in the treatment of oxidative stress using the network pharmacology strategy. The results demonstrated that the majority of bioactive ingredients showed great accumulation at the immature stages in both peels and pulps, and the levels of flavonoid glycosides and terpenoid saponins gradually declined as ripening progressed. Notably, the comparative analysis between the full-blown peel and pulp indicated that the peel had higher concentrations of flavonoids, phenols, and alkaloids, whereas terpenoids and amino acids were the significantly enriched bioactive ingredients in the pulp. In addition, network pharmacology analysis screened 28 key pharmacological compounds in Luohan Guo, which primarily acted on 65 core targets to alleviate diseases induced by oxidative damage and predominantly accumulated in the peels collected between the 10th and 20th days, endowing immature peel with the best antioxidant properties. These findings offer valuabley theoretical insights for understanding the secondary metabolite accumulation in different ripening stages and edible parts of Luohan Guo and improving its comprehensive utilization rate., 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 © 2025 Elsevier Ltd. All rights reserved.)

In this study, to improve the understanding of aroma characteristics, functional components, and nutritional variations in coriander at different growth stages, metabolomic approaches were employed to evaluate the profiles of volatile and non-volatile metabolites in coriander microgreens (CM) and coriander baby greens (CBG), focusing on their types and quantities. A comprehensive analysis identified 1,000 volatile and 1,799 non-volatile metabolites, among which 257 volatile and 597 non-volatile differential metabolites were recognized as key metabolites associated with coriander at various stages of maturity. Notably, key volatile organic compounds (VOCs) 1-p-menthen-8-thiol, 2-isobutyl-3-methoxypyrazine, 5-ethyl-3-hydroxy-4-methyl-2(5H)-furanone and β-ionone were identified as the primary odor determinants during the CM and CBM stages. Additionally, flavonoids such as kaempferol, quercetin, myricetin, isorhamnetin-3-O-glucoside, 7,4'-dihydroxyflavone significantly contribute to flavonoid biosynthesis pathways of coriander. Subsequently, a network pharmacology analysis was conducted to explore the potential therapeutic targets and pathways associated with the bioactive constituents pertinent to type 2 diabetes mellitus (T2DM) and obesity (OB), as identified through the metabolomics analysis. Through comprehensive bioactive ingredient screening, target prediction, protein-protein interaction network analysis, biological process examination, signaling pathway elucidation, and molecular docking validation, the underlying mechanisms by which CM and CBG potentially confer protection against T2DM and OB were elucidated. Through network pharmacology analysis, 10 core targets and their corresponding 49 compounds, including 17 flavonoids, were screened. This study establishes a theoretical framework for the application of CM and CBG as plant-based functional foods, distinguished by their flavor profiles, nutritional content, and health-promoting properties., 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 © 2025 Elsevier Ltd. All rights reserved.)

Rhododendron species have the potential to be rich in secondary metabolites with pharmaceutical or industrial value. However, there is a lack of comprehensive metabolome studies at the genome level, particularly for unique and rare species like Rhododendron bailiense, which exclusively grows in karst environments in Guizhou, southwest China. Recently, genome assembly data for this species was available. In this study, nontargeted metabolomics was employed to investigate the secondary metabolites profile of R. bailiense callus. The callus of R. bailiense was induced using 0.2 mg L -1 TDZ (Thidiazuron) + 0.1 mg L -1 IBA (3-Indole butyric acid). A comparison between light-treated calli and dark-cultured calli revealed differential accumulation of metabolites, particularly in flavonoids, terpenoids, coumarins, and hydroxycinnamic acids, known for their beneficial effects such as antioxidant, anticancer, and anti-inflammatory properties. Proanthocyanidins, with various health-promoting effects, were found to accumulate significantly in dark-cultured calli. Light conditions promoted diterpene and triterpene products, whereas darkness favored sesquiterpene products. Additionally, the study demonstrated the potential of utilizing Agrobacterium transformation technology on callus suspension cells to enhance secondary metabolite production. Comparison with the genome of Rhododendron molle revealed that the R. bailiense genome exhibited active 'glycosyltransferase activity,' possessed a higher number of copies of monoterpene and sesquiterpene terpene synthases, and contained high copies of specific cytochrome P450 members (CYP71, CYP76, CYP79, CYP82, CYP736). This study offers valuable insights and potential strategies for the biosynthesis and production of Rhododendron secondary metabolites with pharmaceutical or industrial significance., (© 2025 Society for Experimental Biology and John Wiley & Sons Ltd.)

Global metabolic and transcriptional reprogramming is a common event in plant abiotic stress responses, however, the relevant molecular mechanisms remain largely unknown. Here, we characterized the physiological function and molecular mechanism for the rice UGT706F1. We found that UGT706F1 can be potently induced by high temperature. Its overexpression can markedly enhance the heat tolerance of rice through improving the capacity of scavenging reactive oxygen species, whereas its functional deletion results in heat sensitivity in rice. To investigate the regulatory mechanism of UGT706F1 in response to high temperature, we carried out extensive screening of the in vitro enzymatic activity of UGT706F1 and discovered that UGT706F1 exhibits broad-spectrum activity toward flavonoid compounds. Through targeted flavonoid metabolomics analysis, we further revealed that the overexpression of UGT706F1 elevated the content of diverse flavonoids and flavonoid glycosides in rice. Subsequently, via transcriptome analysis, we found that following heat treatment, the overexpression of UGT706F1 was capable of enhancing the transcriptional activity of those genes including the flavonoid synthases, heat shock factors, heat shock proteins, glutathione S-transferase, and various antioxidant enzymes. Furthermore, we identified an R2R3 MYB-type transcription factor MYB61 and demonstrated that MYB61 could directly bind the promoter of UGT706F1 and activate the transcription of UGT706F1. The overexpression of MYB61 also enhanced the heat tolerance and increased flavonoid glycosides. Overall, this study unveiled a novel pathway of the plant heat tolerance response mediated by MYB61-UGT706F1 module and identified a new UGT player for the metabolic and transcriptional regulation under high-temperature circumstance., (© 2025 Society for Experimental Biology and John Wiley & Sons Ltd.)

Obesity, often caused by disorders of lipid metabolism, is a global health concern. Flavonoids from staple grains and legumes are expected as a safer and more cost-effective alternative for the future development of dietary flavonoid-based anti-obesity dietary supplements or drugs. This review systematically summarized their content variation, metabolism in the human body, effects and molecular mechanisms on lipid metabolism. These flavonoids intervene in lipid metabolism by inhibiting lipogenesis, promoting lipolysis, enhancing energy metabolism, reducing appetite, suppressing inflammation, enhancing insulin sensitivity, and improving the composition of the gut microbial. Fermentation and sprouting techniques enhance flavonoid content and these beneficial effects. The multidirectional intervention of lipid metabolism is mainly through regulating AMPK signaling pathway. This study provides potential improvement for challenges of application, including addressing high extraction costs and improving bioavailability, ensuring safety, filling clinical study gaps, and investigating potential synergistic effects between flavonoids in grains and legumes, and other components., 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 © 2024 Elsevier Ltd. All rights reserved.)

Blanching is an agricultural practice where vegetables are cultivated in darkness to prevent photosynthesis, thereby modifying their colour, texture, and flavor. The technique is popularly employed in Chinese chive (CC). Blanched Chinese chive (BCC) is renowned for its pale-yellow appearance, delicate flavor, and culinary-medicinal values; nonetheless, how blanching alters the chemical composition largely remains intangible. In this study, the physiological, nutritional, and metabolic profiles of BCC and CC were investigated. In BCC, the contents of ascorbic acid, flavonoids, anthocyanins, etc. were decreased markedly, whereas the sugar content and pungency were increased significantly, indicating that blanching shaped the vegetable flavor. UPLC-ESI-MS/MS analysis revealed 366 differential metabolites between BCC and CC, and the metabolism of flavor precursors, S-alk(en)yl cysteine sulfoxides, were stimulated by blanching. Together, these findings give a strong clue that blanching increases the pungency flavor in CC, and is a useful technique for other selected vegetables., 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 © 2024 Elsevier Ltd. All rights reserved.)

The pharmacological properties of the dried root of Scutellaria baicalensis Georgi, a Chinese medicinal herb, include antioxidant, antibacterial, and antiviral effects. In S. baicalensis quality assessment, concentrations of baicalin, wogonoside, baicalein, and wogonin in the root are crucial. Drought stress commonly affects the biomass and build-up of active compounds in medicinal sections of medicinal plants and thus their quality. The molecular mechanisms underlying the response of S. baicalensis to drought stress remain unexplored. To delve into the impacts of drought stress on the growth and metabolic processes of S. baicalensis, as well as to unravel the underlying molecular mechanisms. We found prolonged and intensified drought treatment causes an initial surge in its fresh weight, plant height, and stem diameter followed by a gradual slowdown, while malondialdehyde (MDA) content rises; while the fresh weight, length, superoxide dismutase (SOD), and catalase (CAT) activities peak before declining, and the root's diameter continuously narrows. In this study, flavonoid index ingredient levels in S. baicalensis initially decreased, then rose as the drought duration extended, followed by a notable post-rehydration increase in baicalin, wogonoside, and baicalein content and decrease in levels of wogonin and oroxylin A. Transcriptome sequencing and KEGG analysis revealed a significant enrichment of DEGs involved in phenylpropanoid biosynthesis and plant hormone signal transduction pathways. The expression levels of SbPAL, SbCCL, Sb4CL, SbCHI, SbFNSII, SbF6H, and SbUGT genes in the flavonoid biosynthetic pathway and PYR/PYL, PP2C, ABF, and SnRK2 genes in the abscisic acid signal transduction pathway were significantly changed. Drought responsive SbWRKY34 was selected for the subsequent investigation. SbWRKY34 showed the highest level in stems, and the encoding protein was localized in the nucleus. Overexpression of SbWRKY34 in Arabidopsis thaliana (OE-SbWRKY34 lines) resulted in increased sensitivity to drought stress, with considerably reduced MDA content and elevated SOD and CAT activities. Concurrently, the expression levels of AtCAT3, AtDREB, AtRD22, AtRD29A, and AtRD29B were significantly reduced in these lines, suggesting that SbWRKY34 functions to negatively regulate drought resistance in A. thaliana., Competing Interests: Declaration of competing interest The authors declare no conflict of interest related to this study., (Copyright © 2024 Elsevier Masson SAS. All rights reserved.)