{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T14:34:20Z","timestamp":1787150060382,"version":"build-2736575974"},"reference-count":48,"publisher":"MDPI AG","issue":"16","license":[{"start":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T00:00:00Z","timestamp":1787097600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100008006","name":"Yibin University program","doi-asserted-by":"publisher","award":["2021QH030"],"award-info":[{"award-number":["2021QH030"]}],"id":[{"id":"10.13039\/501100008006","id-type":"DOI","asserted-by":"publisher"}]},{"award":["2021QH030"],"award-info":[{"award-number":["2021QH030"]}],"id":[{"id":"https:\/\/ror.org\/03w8m2977","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Sichuan Science and Technology Program","award":["2024NSFSC0394"],"award-info":[{"award-number":["2024NSFSC0394"]}]},{"name":"Sichuan Science and Technology Program","award":["2026JDHJ0028"],"award-info":[{"award-number":["2026JDHJ0028"]}]},{"name":"Central Government\u2019s Guiding Funds for Local Scientific and Technological Development","award":["2024ZYD0081"],"award-info":[{"award-number":["2024ZYD0081"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>The flavor variation among different tea plant cultivars significantly influences their economic value. \u2018Huangjinya\u2019 (HJY), a typical light-sensitive etiolated tea cultivar, is prized for its fresh and mellow taste with low bitterness and astringency, and commands a high market value, serving as excellent germplasm for breeding superior-flavor tea varieties. Previous studies have largely focused on the mechanisms underlying fresh and umami tastes, whereas the regulatory mechanisms underlying bitterness and astringency remain poorly understood. In this study, two tea plant (Camellia sinensis (L.) O. Kuntze) cultivars with distinct bitterness and astringency phenotypes, namely \u2018Fuxuan 9\u2019 (FX) and HJY, were used to investigate the formation mechanism of reduced bitterness and astringency through integrated metabolomic and transcriptomic analyses. Metabolomic profiling identified 601 differentially accumulated metabolites between the two cultivars, which were significantly enriched in the flavonoid biosynthesis pathway. Notably, although the total flavonoid content was higher in HJY, the accumulation of non-esterified catechins (C, EC, EGC), which are key contributors to bitterness, was significantly lower than in FX. By contrast, the levels of esterified catechins (EGCG, GCG) showed no marked differences between cultivars. This distinct accumulation pattern provides a metabolic basis potentially associated with the reduced bitterness and astringency in HJY. Transcriptomic analysis revealed that the catechin biosynthetic gene CsLAR was significantly down-regulated in HJY, consistent with the reduced accumulation of non-esterified catechins. Promoter cloning and functional validation confirmed that the CsLAR promoters in both cultivars were active, which differed by only one regulatory element. Under shading treatment, CsLAR expression in HJY was more sensitive to light and decreased more markedly, suggesting that the expression of CsLAR may be regulated by transcription factors. Furthermore, 385 differentially expressed transcription factors were identified, with predicted binding sites in the CsLAR promoter region, implying their potential involvement in modulating catechin accumulation through modulation of CsLAR expression. Together, these multi-omics findings reveal molecular features potentially contributing to the low bitterness and astringency of HJY, providing a theoretical basis and candidate regulators for breeding tea cultivars with improved flavor profiles.<\/jats:p>","DOI":"10.3390\/plants15162508","type":"journal-article","created":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T14:07:34Z","timestamp":1787148454000},"page":"2508","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Integrated Transcriptomics and Metabolomics Analysis of the Formation Mechanism of Low Bitterness and Astringency in \u201cHuangjinya\u201d Tea"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1118-7021","authenticated-orcid":false,"given":"Linghui","family":"Wang","sequence":"first","affiliation":[{"name":"Faculty of Agriculture, Forestry and Food Engineering, Yibin University, Yibin 644001, China"},{"name":"Tea Resources Utilization and Quality Testing Key Laboratory of Sichuan Province, Chengdu 611130, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Quan","family":"Xu","sequence":"additional","affiliation":[{"name":"Faculty of Agriculture, Forestry and Food Engineering, Yibin University, Yibin 644001, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Miao","family":"Xu","sequence":"additional","affiliation":[{"name":"Faculty of Agriculture, Forestry and Food Engineering, Yibin University, Yibin 644001, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xin","family":"Tong","sequence":"additional","affiliation":[{"name":"Faculty of Agriculture, Forestry and Food Engineering, Yibin University, Yibin 644001, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lianxin","family":"Liu","sequence":"additional","affiliation":[{"name":"Faculty of Agriculture, Forestry and Food Engineering, Yibin University, Yibin 644001, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ding","family":"Tang","sequence":"additional","affiliation":[{"name":"Faculty of Agriculture, Forestry and Food Engineering, Yibin University, Yibin 644001, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Suyu","family":"Xia","sequence":"additional","affiliation":[{"name":"Faculty of Agriculture, Forestry and Food Engineering, Yibin University, Yibin 644001, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ji","family":"Huang","sequence":"additional","affiliation":[{"name":"Faculty of Agriculture, Forestry and Food Engineering, Yibin University, Yibin 644001, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei","family":"Li","sequence":"additional","affiliation":[{"name":"Faculty of Agriculture, Forestry and Food Engineering, Yibin University, Yibin 644001, China"},{"name":"Tea Resources Utilization and Quality Testing Key Laboratory of Sichuan Province, Chengdu 611130, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8019-3639","authenticated-orcid":false,"given":"Xia","family":"Zhao","sequence":"additional","affiliation":[{"name":"Faculty of Agriculture, Forestry and Food Engineering, Yibin University, Yibin 644001, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qiulan","family":"Huang","sequence":"additional","affiliation":[{"name":"Faculty of Agriculture, Forestry and Food Engineering, Yibin University, Yibin 644001, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,19]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1016\/j.ygeno.2020.12.020","article-title":"Multiple-genotypes transcriptional analysis revealed candidates genes and nucleotide variants for improvement of quality characteristics in tea (Camellia sinensis (L.) O. Kuntze)","volume":"113","author":"Maritim","year":"2021","journal-title":"Genomics"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"111375","DOI":"10.1016\/j.foodres.2022.111375","article-title":"Comparative transcriptomic analysis reveals the regulatory mechanisms of catechins synthesis in different cultivars of Camellia sinensis","volume":"157","author":"Zhao","year":"2022","journal-title":"Food Res. Int."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Zhang, Q., and Ruan, J. (2016). Tea: Analysis and tasting. Encyclopedia of Food and Health, Academic Press.","DOI":"10.1016\/B978-0-12-384947-2.00687-5"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"107316","DOI":"10.1016\/j.biotechadv.2018.11.005","article-title":"Flavonoid biosynthetic pathways in plants: Versatile targets for metabolic engineering","volume":"38","author":"Nabavi","year":"2020","journal-title":"Biotechnol. Adv."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1039\/C8FO01936C","article-title":"Hypoglycemic potential of whole green tea: Water-soluble green tea polysaccharides combined with green tea extract delays digestibility and intestinal glucose transport of rice starch","volume":"10","author":"Chung","year":"2019","journal-title":"Food Funct."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Furushima, D., Ide, K., and Yamada, H. (2018). Effect of tea catechins on influenza infection and the common cold with a focus on epidemiological\/clinical studies. Molecules, 237.","DOI":"10.3390\/molecules23071795"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1016\/j.tifs.2020.05.015","article-title":"Association between chemistry and taste of tea: A review","volume":"101","author":"Zhang","year":"2020","journal-title":"Trends Food Sci. Technol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1016\/j.tifs.2022.02.031","article-title":"Bitterness and astringency of tea leaves and products: Formation mechanism and reducing strategies","volume":"123","author":"Ye","year":"2022","journal-title":"Trends Food Sci. Technol."},{"key":"ref_9","first-page":"8873091","article-title":"Tea Catechin Inhibits Biofilm Formation of Methicillin-Resistant S. aureus","volume":"20211","author":"Zhao","year":"2021","journal-title":"J. Food Qual."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Chen, X., Wang, P., Zheng, Y., Gu, M., Lin, X., Wang, S., Jin, S., and Ye, N. (2020). Comparison of metabolome and transcriptome of flavonoid biosynthesis pathway in a purple-leaf tea germplasm Jinmingzao and a green-leaf tea germplasm Huangdan reveals their relationship with genetic mechanisms of color formation. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21114167"},{"key":"ref_11","first-page":"E4151","article-title":"Draft genome sequence of Camellia sinensis var. sinensis provides insights into the evolution of the tea genome and tea quality","volume":"11518","author":"Wei","year":"2018","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"2285","DOI":"10.1093\/jxb\/erw053","article-title":"Identification of UDP-glycosyltransferases involved in the biosynthesis of astringent taste compounds in tea (Camellia sinensis)","volume":"678","author":"Cui","year":"2016","journal-title":"J. Exp. Bot."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1016\/j.foodchem.2014.01.044","article-title":"Determination of quality constituents in the young leaves of albino tea cultivars","volume":"155","author":"Feng","year":"2014","journal-title":"Food Chem."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"111569","DOI":"10.1016\/j.plantsci.2022.111569","article-title":"UV-B regulates seasonal greening of albino leaves by modulating CsHY5-inhibiting chlorophyll biosynthesis in Camellia sinensis cv. Huangkui","volume":"328","author":"Liu","year":"2022","journal-title":"Plant Sci."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1016\/j.scienta.2019.03.032","article-title":"Response of leaf color and the expression of photoreceptor genes of Camellia sinensis cv. Huangjinya to different light quality conditions","volume":"251","author":"Tian","year":"2019","journal-title":"Sci. Hortic."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Liu, L., Li, Y., She, G., Zhang, X., Jordan, B., Chen, Q., Zhao, J., and Wan, X. (2018). Metabolite profiling and transcriptomic analyses reveal an essential role of UVR8-mediated signal transduction pathway in regulating flavonoid biosynthesis in tea plants (Camellia sinensis) in response to shading. BMC Plant Biol., 18.","DOI":"10.1186\/s12870-018-1440-0"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"612","DOI":"10.1007\/s00344-021-10325-2","article-title":"An RNA-seq analysis reveals differential transcriptional responses to different light qualities in leaf color of Camellia sinensis cv. Huangjinya","volume":"412","author":"Tian","year":"2022","journal-title":"J. Plant Growth Regul."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Zhang, Q., Liu, M., and Ruan, J. (2017). Integrated transcriptome and metabolic analyses reveals novel insights into free amino acid metabolism in Huangjinya tea cultivar. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.00291"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1016\/j.foodchem.2018.10.010","article-title":"Differential accumulation of specialized metabolite L-theanine in green and albino-induced yellow tea (Camellia sinensis) leaves","volume":"276","author":"Cheng","year":"2019","journal-title":"Food Chem."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1501","DOI":"10.1093\/treephys\/tpac016","article-title":"Integrated transcriptome and amino acid profile analyses reveal novel insights into differential accumulation of theanine in green and yellow tea cultivars","volume":"427","author":"Xu","year":"2022","journal-title":"Tree Physiol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"114739","DOI":"10.1016\/j.lwt.2023.114739","article-title":"Quality components identification and formation analysis of tea (Camellia sinensis L.) flower beverages from three cultivars","volume":"181","author":"Wu","year":"2023","journal-title":"LWT"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"138454","DOI":"10.1016\/j.foodchem.2024.138454","article-title":"Foliar application of iron impacts flavonoid glycosylation and promotes flavonoid metabolism in coloured rice","volume":"444","author":"Zhao","year":"2024","journal-title":"Food Chem."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"22904","DOI":"10.3390\/ijms160922904","article-title":"Transcriptome analysis and identification of differentially expressed transcripts of immune-related genes in spleen of gosling and adult goose","volume":"169","author":"Wang","year":"2015","journal-title":"Int. J. Mol. Sci."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1769","DOI":"10.1093\/mp\/sst080","article-title":"A novel integrated method for large-scale detection, identification, and quantification of widely targeted metabolites: Application in the study of rice metabolomics","volume":"6","author":"Chen","year":"2013","journal-title":"Mol. Plant"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"2565","DOI":"10.1038\/s41467-024-46955-9","article-title":"An ARF gene mutation creates flint kernel architecture in dent maize","volume":"15","author":"Wang","year":"2024","journal-title":"Nat. Commun."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"103074","DOI":"10.1016\/j.fochx.2025.103074","article-title":"Metabolic insights into the enhanced umami and sweetness of small-leaf yellow tea from albino cultivars","volume":"31","author":"Li","year":"2025","journal-title":"Food Chem. X"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"uhac269","DOI":"10.1093\/hr\/uhac269","article-title":"Haem Oxygenase 1 is a potential target for creating etiolated\/albino tea plants (Camellia sinensis) with high theanine accumulation","volume":"102","author":"Chen","year":"2023","journal-title":"Hortic. Res."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"868","DOI":"10.1016\/j.cub.2024.01.044","article-title":"Root microbiota of tea plants regulate nitrogen homeostasis and theanine synthesis to influence tea quality","volume":"344","author":"Xin","year":"2024","journal-title":"Curr. Biol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"138133","DOI":"10.1016\/j.foodchem.2023.138133","article-title":"The metabolic mechanism of flavonoid glycosides and their contribution to the flavor evolution of white tea during prolonged withering","volume":"439","author":"Wang","year":"2024","journal-title":"Food Chem."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"115794","DOI":"10.1016\/j.ab.2025.115794","article-title":"Total flavonoid content revised: An overview of past, present, and future determinations in phytochemical analysis","volume":"700","author":"Nicolescu","year":"2025","journal-title":"Anal. Biochem."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"111932","DOI":"10.1016\/j.lwt.2021.111932","article-title":"Determination of total flavonoid content by aluminum chloride assay: A critical evaluation","volume":"150","author":"Shraim","year":"2021","journal-title":"LWT"},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Zhang, Q., Liu, M., and Ruan, J. (2017). Metabolomics analysis reveals the metabolic and functional roles of flavonoids in light-sensitive tea leaves. BMC Plant Biol., 17.","DOI":"10.1186\/s12870-017-1012-8"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Mei, X., Zhang, K., Lin, Y., Su, H., Lin, C., Chen, B., Haijun, Y., and Zhang, L. (2022). Metabolic and transcriptomic profiling reveals etiolated mechanism in huangyu tea (Camellia Sinensis) leaves. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms232315044"},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Favari, C., Mena, P., Curti, C., Del Rio, D., and Angelino, D. (2020). Flavan-3-ols: Catechins and proanthocyanidins. Dietary Polyphenols: Their Metabolism and Health Effects, Wiley.","DOI":"10.1002\/9781119563754.ch8"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"620","DOI":"10.1016\/j.bbrc.2011.01.079","article-title":"Evaluation of the bitterness of green tea catechins by a cell-Based assay with the human bitter taste receptor hTAS2R39","volume":"405","author":"Narukawa","year":"2011","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1984","DOI":"10.1016\/j.foodhyd.2009.03.001","article-title":"Astringency of tea catechins: More than an oral lubrication tactile percept","volume":"23","author":"Rossetti","year":"2009","journal-title":"Food Hydrocoll."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1016\/j.foodchem.2018.03.042","article-title":"Quantitative analyses of the bitterness and astringency of catechins from green tea","volume":"258","author":"Xu","year":"2018","journal-title":"Food Chem."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"e70249","DOI":"10.1111\/1541-4337.70249","article-title":"Flavone glycosides of tea: Structure, isolation, identification, health benefit and sensory property","volume":"245","author":"Ke","year":"2025","journal-title":"Compr. Rev. Food Sci. Food Saf."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"e71277","DOI":"10.1002\/fsn3.71277","article-title":"Biosynthesis and application of catechins and their derivatives in Camellia sinensis","volume":"1312","author":"Yao","year":"2025","journal-title":"Food Sci. Nutr."},{"key":"ref_40","first-page":"105","article-title":"Pharmacokinetic and chemoprevention studies on tea in humans","volume":"642","author":"Chow","year":"2011","journal-title":"Pharmacol. Res."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"4281","DOI":"10.1021\/acs.jafc.8b00946","article-title":"An insight into catechins metabolic pathways of Camellia sinensis based on genome and transcriptome analysis","volume":"66","author":"Wang","year":"2018","journal-title":"J. Agric. Food Chem."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Lin, J., Lai, G., Guo, A., He, L., Yang, F., Huang, Y., Che, J., and Lai, C. (2024). Overexpression of LAR1 suppresses anthocyanin biosynthesis by enhancing catechin competition leading to promotion of proanthocyanidin pathway in spine grape (Vitis davidii) cells. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms252212087"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"882","DOI":"10.15389\/agrobiology.2022.5.882eng","article-title":"Genetic mechanisms of the biosynthesis of catechins, caffeine and L-theanine in the tea plant Camellia sinensis (L.) Kuntze","volume":"57","author":"Malyukova","year":"2022","journal-title":"Sel\u2019skokhozyaistvennaya Biol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1007\/s11103-005-2910-y","article-title":"The MYB transcription factor superfamily of Arabidopsis: Expression analysis and phylogenetic comparison with the rice MYB family","volume":"60","author":"Yanhui","year":"2006","journal-title":"Plant Mol. Biol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"5441","DOI":"10.1093\/jxb\/erad248","article-title":"Advances in cis-element-and natural variation-mediated transcriptional regulation and applications in gene editing of major crops","volume":"74","author":"Cui","year":"2023","journal-title":"J. Exp. Bot."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Song, X., Huang, X., Li, Q., Lin, H., Bai, S., Zhu, M., Li, J., and Wang, K. (2023). The WRKY transcription factor CsWRKY70 regulates EGCG Biosynthesis by affecting CsLAR and CsUGT84A expressions in Tea leaves (Camellia sinensis). Horticulturae, 9.","DOI":"10.3390\/horticulturae9010120"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1104","DOI":"10.1007\/s00344-024-11281-3","article-title":"Functionality of the MYB1 transcription factor in the transactivation of leucoanthocyanidin reductase (LAR) promoters of Fragaria \u00d7 Ananassa and Fragaria Chiloensis","volume":"44","author":"Figueroa","year":"2025","journal-title":"J. Plant Growth Regul."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"113798","DOI":"10.1016\/j.scienta.2024.113798","article-title":"The R2R3 MYB700 activates the expression of biosynthetic genes involved in the accumulation of (+)-catechin but not (-)-epicatechin in Chinese plum fruits (Prunus salicina Lindl.)","volume":"338","author":"Battistoni","year":"2024","journal-title":"Sci. Hortic."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/16\/2508\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T14:15:10Z","timestamp":1787148910000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/16\/2508"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,19]]},"references-count":48,"journal-issue":{"issue":"16","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["plants15162508"],"URL":"https:\/\/doi.org\/10.3390\/plants15162508","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,19]]}}}