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Saito, K., Yonekura-Sakakibara, K., Nakabayashi, R., Higashi, Y., Yamazaki, M., Tohge, T., et al. (2013). The flavonoid biosynthetic pathway in arabidopsis: structural and genetic diversity. Plant Physiol. Biochem. 72, 21–34. doi:10.1016/j.plaphy.2013.02.001 Forkmann, G., Martens, S. (2001). Metabolic engineering and applications of flavonoids. Curr. Opin. Biotechnol. 12, 155–160. doi:10.1016/s0958-1669(00)00192-0 CiT Pro Android X Gaming Cube White Case with 3 x 120mm Infinity ARGB Fans 1 x 6-Port Fan Hub Tempered Glass Front and Side Panels This study was financially supported by the National Key R&D Program of China (2021YFD1600802-02), the Science and Rechnology Department of Sichuan Province, China (2021ZHCG0084), the 14th- fifth-plan of Breeding in Sichuan Province, China (2021YFYZ0023-14). Conflict of interest Owens, D. K., Alerding, A. B., Crosby, K. C., Bandara, A. B., Westwood, J. H., Winkel, B. S. (2008). Functional analysis of a predicted flavonol synthase gene family in arabidopsis. Plant Physiol. 147, 1046–1061. doi:10.1104/pp.108.117457
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Li, Y., Chen, Q. Y., Xie, X. D., Cai, Y., Li, J. F., Feng, Y. L., et al. (2020b). Integrated metabolomics and transcriptomics analyses reveal the molecular mechanisms underlying the accumulation of anthocyanins and other flavonoids in cowpea pod (Vigna unguiculata l.). J. Agric. Food Chem. 68, 9260–9275. doi:10.1021/acs.jafc.0c01851 Hu, H., Fei, X., He, B., Luo, Y., Qi, Y., Wei, A. (2021). Integrated analysis of metabolome and transcriptome data for uncovering flavonoid components of zanthoxylum bungeanum maxim. leaves under drought stress. Front. Nutr. 8. doi:10.3389/fnut.2021.801244 Espley, R. V., Hellens, R. P., Putterill, J., Stevenson, D. E., Kutty-Amma, S., Allan, A. C. (2007). Red colouration in apple fruit is due to the activity of the MYB transcription factor, MdMYB10. Plant J. 49, 414–427. doi:10.1111/j.1365-313X.2006.02964.x Alseekh, S., Aharoni, A., Brotman, Y., Contrepois, K., D’auria, J., Ewald, J., et al. (2021). Mass spectrometry-based metabolomics: a guide for annotation, quantification and best reporting practices. Nat. Methods 18, 747–756. doi:10.1038/s41592-021-01197-1
Introduction
Khare, S., Singh, N. B., Singh, A., Hussain, I., Niharika, K., Yadav, V., et al. (2020). Plant secondary metabolites synthesis and their regulations under biotic and abiotic constraints. J. Plant Biol. 63, 203–216. doi:10.1007/s12374-020-09245-7 Mahmoud, A. M., Hernandez Bautista, R. J., Sandhu, M. A., Hussein, O. E. (2019). Beneficial effects of citrus flavonoids on cardiovascular and metabolic health. Oxid. Med. Cell Longev 2019, 5484138. doi:10.1155/2019/5484138 In GO annotation analysis, a total of 38,397 DEGs were annotated in three categories: biological process, molecular function, and cellular component categories ( Supplementary Table7). These DEGs were further divided into 47 categories based on gene function, with 722 genes related to biological processes such as signaling. TopGO analysis revealed that the most enriched molecular function terms were monooxygenase activity (GO0004497), oxidoreductase activity (GO0016705), heme binding (GO0020037), iron ion binding (GO0005506), and tetrapyrrole binding (GO0046906) ( Supplementary Figure7). The most enriched biological process terms were flavonoid metabolic process (GO0009812) and flavonoid biosynthetic process (GO0009813). The most enriched cellular component terms were chloroplast thylakoid (GO0009534) and plastid thylakoid (GO0031976). KEGG enrichment analysis identified the top 20 enriched metabolic pathways, including metabolic pathways (ko01100), biosynthesis of secondary metabolites (ko01110), MAPK signaling pathway-plant (ko04016), plant hormone signal transduction (ko04075), phenylpropanoid biosynthesis (ko00940), and flavonoid biosynthesis (ko00941) under magnesium stress ( Figure5D). These results were presented in a bubble diagram. Metabolic and gene co-expression networks in SOPs at different developmental stages Wang, Y., Liu, X. J., Chen, J. B., Cao, J. P., Li, X., Sun, C. D. (2022). Citrus flavonoids and their antioxidant evaluation. Crit. Rev. Food Sci. Nutr. 62, 3833–3854. doi:10.1080/10408398.2020.1870035 An improved protocol was used to determine the total flavonoid content of citrus peels ( Wang etal., 2007; Yu etal., 2022). Initially, 0.5 g citrus peel powder was weighed and dissolved in 10 mL 70% absolute ethanol at a ratio of 1:20 (w/v). The mixture was then subjected to ultrasonic treatment at 55°C for 40 min, followed by filtration. To 1 mL of the extraction solution, 0.5 mL of 5% NaNO 2 solution was added sequentially and well shaken. The mixture was then left for 5 min. Next, 0.5 mL 10% Al(NO 3) 3 was added to the solution, mixed thoroughly, and left for 6 min. Finally, 5 mL of 1mol/L NaOH was added, and distilled water was added up to 10 mL. The mixture was shaken and left for 10 min to complete the reaction. The absorbance value of solution was measured at 510 nm, with rutin (purity≥98%, sourced from Leaf Shanghai Biological Technology Co., Ltd.) used as the standard product. The flavonoid content (U mol/g) was calculated using the formula (C*V)/W, where C represents the concentration, V represents the volume, and W represents the weight of the sample. To determine the MDA content and SOD activity, Li’s method was followed ( Li etal., 2022b). Metabolomic profile detection and analysis
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Lepiniec, L., Debeaujon, I., Routaboul, J. M., Baudry, A., Pourcel, L., Nesi, N., et al. (2006). Genetics and biochemistry of seed flavonoids. Annu. Rev. Plant Biol. 57, 405–430. doi:10.1146/annurev.arplant.57.032905.105252 Citation: Xiong B, Li Q, Yao J, Liu Z, Yang X, Yu X, Li Y, Liao L, Wang X, Deng H, Zhang M, Sun G and Wang Z (2023) Widely targeted metabolomic profiling combined with transcriptome analysis sheds light on flavonoid biosynthesis in sweet orange 'Newhall' (C. sinensis) under magnesium stress. Front. Plant Sci. 14:1182284. doi: 10.3389/fpls.2023.1182284 Project and experiment design, ZW and BX. Experiment execution, QL, JY, ZL, XXY, and XYY. Data analysis, QL, YL, LL, XW, HD, and MZ. Writing, BX and QL. Review, BX and ZW. Project management, BX, GS, and ZW. All authors contributed to the article and approved the submitted version. Funding Hichri, I., Barrieu, F., Bogs, J., Kappel, C., Delrot, S., Lauvergeat, V. (2011). Recent advances in the transcriptional regulation of the flavonoid biosynthetic pathway. J. Exp. Bot. 62, 2465–2483. doi:10.1093/jxb/erq442 Deng, Y. X., Lu, S. F. (2017). Biosynthesis and regulation of phenylpropanoids in plants. Crit. Rev. Plant Sci. 36, 257–290. doi:10.1080/07352689.2017.1402852
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Bartwal, A., Mall, R., Lohani, P., Guru, S. K., Arora, S. (2013). Role of secondary metabolites and brassinosteroids in plant defense against environmental stresses. J. Plant Growth Regul. 32, 216–232. doi:10.1007/s00344-012-9272-x The first two principal components accounted for 50.35% (PC1) and 17.79% (PC2), respectively, and the 18 samples (including 3 replicates) were classified into 6 groups based on their developmental stage along PC1. The sample positions along PC2 were influenced by magnesium stress ( Figure3B). These findings suggest that the observed differences in flavonoid profiles were related to developmental stages and magnesium stress and were consistent with the trend in total flavonoid accumulation that peaked in MS2 or MD2 ( Figures3B, C). In addition, OPLS-DA analysis was utilized to evaluate the differences between MS and MD (Q2 = 0.99) ( Supplementary Figure2). The high Q2 value (>0.9) suggested that the OPLS-DA modules were stable and reliable and that the differences in flavonoid content could be further explored. Hierarchical clustering analysis (HCA) of the flavonoid metabolite accumulation patterns among different samples showed good repeatability within the sample group ( Figures3D). In the HCA, six clusters, corresponding to the successive stages of flavonoid metabolites in SOPs for the 18 samples, were significantly separated. The results of PCA, OPLS-DA, correlation analysis, and HCA reflected large differences between samples, high similarity among the three biological replicates, and high repeatability within samples. Differentially accumulated flavonoids metabolites in SOPs An investigation of differentially accumulated flavonoids (DAFs) was conducted in SOPs at different development stages. A total of 740 flavonoids were screened, and 142 DAFs were selected based on a fold change of |log2FC| ≥ 2 or |log2FC| ≤ 0.5 and a variable importance in projection (VIP ≥1) ( Supplementary Table4). Of these, 57 DAFs were identified in MS1 vs. MD1, followed by MS2 vs. MD2 (25) and MS3 vs. MD3 (97) ( Figure4C). The Venn Diagram results revealed two common and unique differential metabolites (Chrysoeriol-7-O-glucoside, Chrysoeriol-7-O-(6’’-feruloyl) glucoside) between MS and MD across all three periods. These flavonoids were flavones, and their change trend was consistent with total flavonoid content. To study the variation of these differential metabolites under magnesium stress, volcano diagrams were performed ( Supplementary Figure5). The results indicated that there were more up-regulated than down-regulated flavonoids in three stages between MS and MD. Specifically, 57 DAFs (53 upregulated and 4 downregulated) were identified during MS1 vs. MD1, 25 DAFs (6 upregulated and 19 downregulated) were identified during MS2 vs. MD2, and 97 DAFs (86 upregulated and 11 downregulated) were identified during MS3 vs. MD3. The majority of DAFs were observed during the development period. There were 278 DAFs (141 upregulated and 137 downregulated) and 261 DAFs (161 upregulated and 100 downregulated) selected from MD1 vs. MD2 and MS1 vs. MS2, respectively. The greater number of DAFs in MD than MS suggested that flavonoids may have been more susceptible to magnesium stress. The interaction of DAFs in SOPs resulted in the formation of different pathways, which were annotated and assigned to the KEGG pathways ( Figure4D). KEGG pathway enrichment analysis showed that flavonoid biosynthesis, phenylpropanoid biosynthesis, flavone and flavonol biosynthesis, secondary metabolites biosynthesis and metabolic pathways were the main enrichment pathways. Therefore, it could be postulated that the differentially accumulated metabolites (DAMs) in the pathways mentioned above may contribute to the variation in flavonoids of SOPs during the developmental process. Differentially expressed gene analysis Shen, N., Wang, T., Gan, Q., Liu, S., Wang, L., Jin, B. (2022). Plant flavonoids: classification, distribution, biosynthesis, and antioxidant activity. Food Chem. 383, 132531. doi:10.1016/j.foodchem.2022.132531
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CiT Vento White Micro-ATX PC Gaming Case with 4 x 120mm ARGB Fans Included 1 x 6-Port Fan Hub Tempered Glass Side Panel Increased Air Flow - Top & Front panel incorporate mesh paneling, to improve air flowplastic and mesh steelCiT Level 2 White Micro-ATX Mesh PC Gaming Case with 3 x 120mm RGB Rainbow Fans Included With Tempered Glass Side Panel Ramakrishna, A., Ravishankar, G. A. (2011). Influence of abiotic stress signals on secondary metabolites in plants. Plant Signal Behav. 6, 1720–1731. doi:10.4161/psb.6.11.17613 Citrus fruits, members of the Rutaceae family, are widely consumed throughout the world. A prime example is the sweet orange ‘Newhall’ ( C. sinensis), which stands out for its exceptional quality and has its origins in America. Citrus fruits are a rich source of bioactive flavonoids, with the peels often containing a higher concentration of these compounds than pulp and seeds ( Li etal., 2022a; Yu etal., 2022). Previous studies have shown that flavonoid compounds in Citrus peel play a significant role in anti-inflammation, anti-oxidation, immune regulation, and prevention and treatment of multiple respiratory diseases ( Peng etal., 2019; Singh etal., 2020). Unlike most other fruits, Citrus species mainly accumulate flavonone glycosides and polymethoxylated flavones (PMFs) as their main flavonoids ( Zhao etal., 2021). These compounds are highly valued as a source of common Chinese medicines, food, and nutritional supplements due to their abundance of bioactive components.
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Bo Xiong *† Qin Li † Junfei Yao Zhuyuan Liu Xinxia Yang Xiaoyong Yu 1 Yuan Li Ling Liao Xun Wang Honghong Deng Mingfei Zhang Guochao Sun Zhihui Wang * Nair, S. A., Kurup, S. R. R., Nair, A. S., Baby, S. (2018). Citrus peels prevent cancer. Phytomedicine 50, 231–237. doi:10.1016/j.phymed.2017.08.011 Environmental factors can regulate gene expression by influencing TFs, which bind specifically to the promoters of their target genes. Among TF families, the MYB family has been shown to play a critical role in regulating gene expression in the flavonoid pathway ( Espley etal., 2007; Zhou etal., 2015; Zhai etal., 2016; Li etal., 2020a). For instance, the MdBBX22–miR858–MdMYB9/11/12 was found to activate the promoters of MdANR and MdLAR in apple, thereby promoting the biosynthesis of proanthocyanidin ( Zhang etal., 2022). In this study, one CitMYB (Cs_ont_1g021030) was identified as highly related to structural genes and seven flavonoids based on WGCNA. Therefore, these ten TF genes were considered important in regulating the flavonoid content of SOPs. Although the results of qRT-PCR showed good consistency with the transcriptome data ( Supplementary Figure10), future studies are needed to elucidate the function of these genes in flavonoid biosynthesis. Conclusion Peng, Y., Hu, M. J., Lu, Q., Tian, Y., He, W. Y., Chen, L., et al. (2019). Flavonoids derived from exocarpium citri grandis inhibit LPS-induced inflammatory response via suppressing MAPK and NF-kappa b signalling pathways. Food Agric. Immunol. 30, 564–580. doi:10.1080/09540105.2018.1550056CiT LT100 1 Litre USB3.0 Ultra-Thin Mini-ITX Computer Case With 120W Laptop Adpater CPU Cooler WIFI Antenna 5cm Included Zhao, C. N., Liu, X. J., Gong, Q., Cao, J. P., Shen, W. X., Yin, X. R., et al. (2021). Three AP2/ERF family members modulate flavonoid synthesis by regulating type IV chalcone isomerase in citrus. Plant Biotechnol. J. 19, 671–688. doi:10.1111/pbi.13494 Li, H., Li, Y., Yu, J. X., Wu, T., Zhang, J., Tian, J., et al. (2020a). MdMYB8 is associated with flavonol biosynthesis via the activation of the MdFLS promoter in the fruits of malus crabapple. Horticult. Res. 7. doi:10.1038/s41438-020-0238-z Song, Z., Luo, Y., Wang, W., Fan, N., Wang, D., Yang, C., et al. (2019). NtMYB12 positively regulates flavonol biosynthesis and enhances tolerance to low pi stress in nicotiana tabacum. Front. Plant Sci. 10. doi:10.3389/fpls.2019.01683 Treutter, D. (2005). Significance of flavonoids in plant resistance and enhancement of their biosynthesis. Plant Biol. (Stuttg) 7, 581–591. doi:10.1055/s-2005-873009
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