{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T14:09:03Z","timestamp":1779199743579,"version":"3.51.4"},"reference-count":56,"publisher":"MDPI AG","issue":"5","license":[{"start":{"date-parts":[[2026,4,26]],"date-time":"2026-04-26T00:00:00Z","timestamp":1777161600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"National Modern Agricultural Industry Technology System","award":["CARS-21"],"award-info":[{"award-number":["CARS-21"]}]},{"name":"Henan Provincial Modern Agricultural Industry Technology System","award":["HARS-22-11-G3"],"award-info":[{"award-number":["HARS-22-11-G3"]}]},{"DOI":"10.13039\/501100017700","name":"Henan Provincial Science and Technology Research Project","doi-asserted-by":"crossref","award":["252102110151"],"award-info":[{"award-number":["252102110151"]}],"id":[{"id":"10.13039\/501100017700","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Genes"],"abstract":"<jats:p>Background: Pinellia ternata is a major medicinal herb widely utilized in traditional medicine, but is sensitive to high temperature, which often triggers a severe \u201csprout tumble\u201d phenomenon. Methods: To elucidate the molecular mechanisms of heat tolerance in P. ternata, we screened two contrasting germplasms: the heat-tolerant JBX1 and the heat-sensitive XBX4. In the present study, a combined analysis of physiology, transcriptome, and metabolome was performed on JBX1 and XBX4 under heat stress at 40 \u00b0C. Results: JBX1 exhibited significantly greater leaf thickness, higher basal chlorophyll content, more stable antioxidant enzyme activities, and lower oxidative damage than XBX4 under heat stress. Transcriptomically, JBX1 maintained elevated basal expression of genes encoding key enzymes in carbon fixation, amino acid metabolism, and phenylpropanoid biosynthesis, as well as those encoding heat shock transcription factors (HSFs), heat shock proteins (HSPs), and the thermosensor Thermo-With ABA-Response 1 (TWA1). Metabolomically, JBX1 accumulated higher levels of key primary metabolites, antioxidants, and protective phenylpropanoids under both control and heat conditions. Notably, a \u201cpolarity reversal\u201d emerged in nitrogen metabolism, where core amino acids accumulated in JBX1 but were depleted in XBX4. Integrated analysis revealed a more coordinated gene\u2013metabolite network in JBX1 involving the phenylpropanoid, ATP-binding cassette (ABC) transporter, and glutathione pathways. Conclusions: Our findings demonstrate that JBX1 possessed stronger basal thermotolerance, which is derived from coordinated establishment of higher constitutive metabolic reserves and efficient dynamic metabolic reprogramming. This study provides insights into the molecular mechanisms of heat stress in P. ternata.<\/jats:p>","DOI":"10.3390\/genes17050512","type":"journal-article","created":{"date-parts":[[2026,4,29]],"date-time":"2026-04-29T07:43:51Z","timestamp":1777448631000},"page":"512","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Integrated Metabolomic and Transcriptomic Analyses Reveal the Differential Molecular Mechanisms Underlying Heat Stress Responses in Two Pinellia ternata Germplasms"],"prefix":"10.3390","volume":"17","author":[{"given":"Guixia","family":"Shi","sequence":"first","affiliation":[{"name":"Institute of Chinese Herbal Medicines, Henan Academy of Agricultural Sciences, Zhengzhou 450002, China"},{"name":"Provincial Key Laboratory of Conservation and Utilization of Henan Traditional Chinese Medicine Resources, Zhengzhou 450002, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Zhen","family":"Yang","sequence":"additional","affiliation":[{"name":"Xixian Agricultural Science Research Institute, Xinyang 464300, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Guixiao","family":"La","sequence":"additional","affiliation":[{"name":"Institute of Chinese Herbal Medicines, Henan Academy of Agricultural Sciences, Zhengzhou 450002, China"},{"name":"Provincial Key Laboratory of Conservation and Utilization of Henan Traditional Chinese Medicine Resources, Zhengzhou 450002, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Miao","family":"Huang","sequence":"additional","affiliation":[{"name":"Xixian Agricultural Science Research Institute, Xinyang 464300, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yulong","family":"Zhao","sequence":"additional","affiliation":[{"name":"Institute of Chinese Herbal Medicines, Henan Academy of Agricultural Sciences, Zhengzhou 450002, China"},{"name":"Provincial Key Laboratory of Conservation and Utilization of Henan Traditional Chinese Medicine Resources, Zhengzhou 450002, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yaping","family":"Li","sequence":"additional","affiliation":[{"name":"Institute of Chinese Herbal Medicines, Henan Academy of Agricultural Sciences, Zhengzhou 450002, China"},{"name":"Provincial Key Laboratory of Conservation and Utilization of Henan Traditional Chinese Medicine Resources, Zhengzhou 450002, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Tiegang","family":"Yang","sequence":"additional","affiliation":[{"name":"Institute of Chinese Herbal Medicines, Henan Academy of Agricultural Sciences, Zhengzhou 450002, China"},{"name":"Provincial Key Laboratory of Conservation and Utilization of Henan Traditional Chinese Medicine Resources, Zhengzhou 450002, China"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2026,4,26]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1612","DOI":"10.1016\/j.molp.2023.09.013","article-title":"The molecular basis of heat stress responses in plants","volume":"16","author":"Kan","year":"2023","journal-title":"Mol. Plant"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1016\/j.plaphy.2012.02.016","article-title":"ROS signaling as common element in low oxygen and heat stresses","volume":"59","author":"Pucciariello","year":"2012","journal-title":"Plant Physiol. Biochem."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"117","DOI":"10.3390\/ijms22010117","article-title":"Plant responses to heat stress: Physiology, transcription, noncoding RNAs, and epigenetics","volume":"22","author":"Zhao","year":"2020","journal-title":"Int. J. Mol. Sci."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"113252","DOI":"10.1016\/j.jep.2020.113252","article-title":"Pinellia ternata (Thunb.) Breit: A review of its germplasm resources, genetic diversity and active components","volume":"263","author":"Mao","year":"2020","journal-title":"J. Ethnopharmacol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"147426","DOI":"10.1016\/j.gene.2023.147426","article-title":"Pinellia genus: A systematic review of active ingredients, pharmacological effects and action mechanism, toxicological evaluation, and multi-omics application","volume":"870","author":"Chen","year":"2023","journal-title":"Gene"},{"key":"ref_6","first-page":"1","article-title":"Pinelliae Rhizoma: A systematic review on botany, ethnopharmacology, phytochemistry, preclinical and clinical evidence","volume":"23","author":"Liang","year":"2025","journal-title":"Chin. J. Nat. Med."},{"key":"ref_7","first-page":"2020","article-title":"Programmed cell death induced by drought stress in sprout tumble of Pinellia ternata","volume":"44","author":"Huang","year":"2019","journal-title":"China J. Chin. Mater. Med."},{"key":"ref_8","first-page":"692","article-title":"Study on the Mechanisms and prevention techniques of sprout tumbling in Suzhou Pinellia ternata","volume":"31","author":"Liu","year":"2003","journal-title":"J. Anhui Agri. Sci."},{"key":"ref_9","first-page":"11","article-title":"Effects of different temperatures on sprout tumbling of Pinellia ternata","volume":"6","author":"Ma","year":"2021","journal-title":"Yunnan Agric. Sci. Technol."},{"key":"ref_10","first-page":"15","article-title":"Research progress on Pinellia ternata","volume":"2","author":"Guo","year":"2000","journal-title":"Chin. J. Tradit. Med. Sci. Technol."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Fanourakis, D., Tsaniklidis, G., Makraki, T., Nikoloudakis, N., Bartzanas, T., Sabatino, L., Fatnassi, H., and Ntatsi, G. (2025). Climate change impacts on greenhouse horticulture in the mediterranean basin: Challenges and adaptation strategies. Plants, 14.","DOI":"10.3390\/plants14213390"},{"key":"ref_12","first-page":"532","article-title":"Experimental observation on preventing sprout tumbling of Pinellia ternata by summer mulching with wheat straw (Chaff)","volume":"21","author":"Pan","year":"1996","journal-title":"China J. Chin. Mater. Med."},{"key":"ref_13","first-page":"774","article-title":"Effects of cultivation measures on sprout tumbling rate and yield of Pinellia ternata","volume":"39","author":"Zhang","year":"2011","journal-title":"J. Anhui Agric. Sci."},{"key":"ref_14","first-page":"2489","article-title":"Change of endogenous hormone around sprout tumble of Pinellia ternata under high temperature stress","volume":"32","author":"Xue","year":"2007","journal-title":"China J. Chin. Mater. Med."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Xue, T., Zhang, H., Zhang, Y.Y., Wei, S.Q., Chao, Q.J., Zhu, Y.F., Teng, J.T., Zhang, A.M., Sheng, W., and Duan, Y.B. (2019). Full-length transcriptome analysis of shade induced promotion of tuber production in Pinellia ternata. BMC Plant Biol., 19.","DOI":"10.1186\/s12870-019-2197-9"},{"key":"ref_16","first-page":"390","article-title":"Structure of cell and histology around sprout tumble of Pinellia ternata","volume":"29","author":"Sheng","year":"2009","journal-title":"Guihaia"},{"key":"ref_17","first-page":"641","article-title":"The change of activity of protective enzyme around sprout tumble of Pinellia ternata under high temperatures stress","volume":"29","author":"Xue","year":"2004","journal-title":"Chin. J. Chin. Mater. Med."},{"key":"ref_18","first-page":"2233","article-title":"Changes of photosynthesis parameters and chlorophyll fluorescence around sprout tumble of Pinellia ternata under high temperature stress","volume":"35","author":"Xue","year":"2010","journal-title":"Chin. J. Chin. Mater. Med."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"20614","DOI":"10.3390\/ijms141020614","article-title":"A comparative proteomic analysis of Pinellia ternata leaves exposed to heat stress","volume":"14","author":"Zhu","year":"2013","journal-title":"Int. J. Mol. Sci."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1007\/s10265-023-01446-8","article-title":"Integrated analysis of metabolome and transcriptome reveals key candidate genes involved in flavonoid biosynthesis in Pinellia ternata under heat stress","volume":"136","author":"Guo","year":"2023","journal-title":"J. Plant Res."},{"key":"ref_21","first-page":"243","article-title":"Physiological and transcriptional responses to heat stress in a typical phenotype of Pinellia ternata","volume":"21","author":"Wang","year":"2023","journal-title":"Chin. J. Nat. Med."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Chen, B., Liu, M.M., You, Q., Liu, X., Zhu, Y.F., Duan, Y.B., Wang, D.X., Xue, T., and Xue, J.P. (2024). Integrated analysis of transcriptome and miRNAome reveals the heat stress response of Pinellia ternata seedlings. BMC Genom., 25.","DOI":"10.1186\/s12864-024-10318-x"},{"key":"ref_23","first-page":"341","article-title":"Variation analysis of genomic methylation induced by high temperature stress in Pinellia ternata","volume":"45","author":"Chao","year":"2020","journal-title":"Chin. J. Chin. Mater. Med."},{"key":"ref_24","first-page":"2614","article-title":"Effects of Ca2 on photosynthetic parameters of Pinellia ternata and accumulations of active components in heat stress","volume":"39","author":"Yang","year":"2014","journal-title":"Chin. J. Chin. Mater. Med."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"110877","DOI":"10.1016\/j.ecoenv.2020.110877","article-title":"Transcriptomic analysis of short-term heat stress response in Pinellia ternata provided novel insights into the improved thermotolerance by spermidine and melatonin","volume":"202","author":"Ma","year":"2020","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"108539","DOI":"10.1016\/j.plaphy.2024.108539","article-title":"Comprehensive in silico characterization of NAC transcription factor family of Pinellia ternata and functional analysis of PtNAC66 under high-temperature tolerance in transgenic Arabidopsis thaliana","volume":"208","author":"Chen","year":"2024","journal-title":"Plant Physiol. Biochem."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.plaphy.2022.02.017","article-title":"Functional characterization of the Pinellia ternata cytoplasmic class II small heat shock protein gene PtsHSP17.2 via promoter analysis and overexpression in tobacco","volume":"177","author":"Tian","year":"2022","journal-title":"Plant Physiol. Biochem."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"13807","DOI":"10.1038\/s41598-024-64560-0","article-title":"PtWRKY2, a WRKY transcription factor from Pinellia ternata confers heat tolerance in Arabidopsis","volume":"14","author":"Liu","year":"2023","journal-title":"Sci. Rep."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"113490","DOI":"10.1016\/j.indcrop.2021.113490","article-title":"Stearic acid desaturase gene negatively regulates the thermotolerance of Pinellia ternata by modifying the saturated levels of fatty acids","volume":"166","author":"Zhang","year":"2021","journal-title":"Ind. Crops Prod."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"824","DOI":"10.1016\/j.tibs.2022.05.004","article-title":"How do plants feel the heat and survive?","volume":"47","author":"Guihur","year":"2022","journal-title":"Trends Biochem. Sci."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Chaudhary, S., Devi, P., Bhardwaj, A., Jha, U.C., Sharma, K.D., Prasad, P.V.V., Siddique, K.H.M., Bindumadhava, H., Kumar, S., and Nayyar, H. (2020). Identification and characterization of contrasting genotypes\/cultivars for developing heat tolerance in agricultural crops: Current status and prospects. Front. Plant Sci., 11.","DOI":"10.3389\/fpls.2020.587264"},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Kumar, A., Muthuramalingam, P., Kumar, R., Tiwari, S., Verma, L., Park, S., and Shin, H. (2025). Adapting crops to rising temperatures: Understanding heat stress and plant resilience mechanisms. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms262110426"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1111\/j.1365-3040.2011.02336.x","article-title":"ROS and redox signalling in the response of plants to abiotic stress","volume":"35","author":"Suzuki","year":"2012","journal-title":"Plant Cell Environ."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"823","DOI":"10.1080\/03650340.2012.683424","article-title":"Effect of varying high temperatures during reproductive growth on reproductive function, oxidative stress and seed yield in chickpea genotypes differing in heat sensitivity","volume":"59","author":"Kumar","year":"2013","journal-title":"Arch. Agron. Soil Sci."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1016\/j.scienta.2016.10.033","article-title":"Influence of high temperature stress on growth, phenology and yield performance of mungbean [Vigna radiata (L.) Wilczek] under managed growth conditions","volume":"213","author":"Sharma","year":"2016","journal-title":"Sci. Hortic."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Sita, K., Sehgal, A., Kumar, J., Kumar, S., Singh, S., Siddique, K.H.M., and Nayyar, H. (2017). Identification of high-temperature tolerant lentil (Lens culinaris Medik.) genotypes through leaf and pollen traits. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.00744"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1007\/s11738-019-2805-1","article-title":"Oxidative damage and antioxidant mechanism in tomatoes responding to drought and heat stress","volume":"41","author":"Zhou","year":"2019","journal-title":"Acta Physiol. Plant."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1023\/A:1026730008917","article-title":"Induction of oxidative stress and antioxidant activity by hydrogen peroxide treatment in tolerant and susceptible wheat genotypes","volume":"43","author":"Sairam","year":"2000","journal-title":"Biol. Plant."},{"key":"ref_39","first-page":"110","article-title":"Oxidative stress and antioxidative system in Brassica juncea (L.) under high temperature stress","volume":"28","author":"Rani","year":"2012","journal-title":"Ann. Biol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"603","DOI":"10.1007\/s11103-013-0013-8","article-title":"Plant senescence and crop productivity","volume":"82","author":"Gregersen","year":"2013","journal-title":"Plant Mol. Biol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1007\/s10681-012-0780-3","article-title":"Association of staygreen trait with canopy temperature depression and yield traits under terminal heat stress in wheat (Triticum aestivum L.)","volume":"190","author":"Kumari","year":"2013","journal-title":"Euphytica"},{"key":"ref_42","first-page":"9","article-title":"Screening wheat germplasm for heat tolerance at terminal growth stage","volume":"2","author":"Rehman","year":"2009","journal-title":"Plant Omics"},{"key":"ref_43","first-page":"14","article-title":"Identification of contrasting genotypes under heat stress in maize (Zea mays L.)","volume":"5","author":"Debnath","year":"2016","journal-title":"Maize J."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"921","DOI":"10.1007\/s12298-020-00801-6","article-title":"Characterization of contrasting genotypes reveals general physiological and molecular mechanisms of heat-stress adaptation in maize (Zea mays L.)","volume":"26","author":"Singh","year":"2020","journal-title":"Physiol. Mol. Biol. Plants"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1334","DOI":"10.1071\/FP13082","article-title":"Heat-stress-induced reproductive failures in chickpea (Cicer arietinum) are associated with impaired sucrose metabolism in leaves and anthers","volume":"40","author":"Kaushal","year":"2013","journal-title":"Funct. Plant Biol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1016\/j.jplph.2004.07.014","article-title":"High temperature effects on photosynthetic activity of two tomato cultivars with different heat susceptibility","volume":"162","author":"Camejo","year":"2005","journal-title":"J. Plant Physiol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1016\/j.plaphy.2019.05.031","article-title":"Chlorophyll fluorescence and carbohydrate concentration as field selection traits for heat tolerant chickpea genotypes","volume":"141","author":"Makonya","year":"2019","journal-title":"Plant Physiol. Biochem."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1941","DOI":"10.1046\/j.1365-3040.2003.01110.x","article-title":"Rubisco expression in rice leaves is related to genotypic variation of photosynthesis under elevated growth CO2 and temperature","volume":"26","author":"Gesch","year":"2003","journal-title":"Plant Cell Environ."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1111\/nph.20017","article-title":"Heat stress transcription factors as the central molecular rheostat to optimize plant survival and recovery from heat stress","volume":"244","author":"Bakery","year":"2024","journal-title":"New Phytol."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Zhang, J.L., Zhu, Y.C., Ma, F.M., Zou, X., Lan, Q.X., Zhou, X.R., Li, M.X., Zhou, F., Yin, C.X., and Lin, Y.J. (2025). Molecular mechanisms of the phytohormone-heat shock protein pathway in regulating plant thermotolerance. Plants, 14.","DOI":"10.3390\/plants14233706"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1126","DOI":"10.1038\/s41586-024-07424-x","article-title":"The temperature sensor TWA1 is required for thermotolerance in Arabidopsis","volume":"629","author":"Bohn","year":"2024","journal-title":"Nature"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1104\/pp.106.091322","article-title":"A heat-inducible transcription factor, HsfA2, is required for extension of acquired thermotolerance in Arabidopsis","volume":"143","author":"Charng","year":"2007","journal-title":"Plant Physiol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"604","DOI":"10.1007\/s004250050524","article-title":"Improving the thiobarbituric acid-reactive-substances assay for estimating lipid peroxidation in plant tissues containing anthocyanin and other interfering compounds","volume":"207","author":"Hodges","year":"1999","journal-title":"Planta"},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Zhao, Y.L., Wang, Y.S., Liu, H.H., Jiang, H.F., Xue, R.L., Wang, S.L., Tai, F.J., Bai, X.R., Fan, G.Q., and Hu, X.L. (2025). Chloroplast CASEIN KINASE 2 enhances maize thermotolerance via sHSP26 phosphorylation and retrograde HSFA3 regulation. Plant Physiol., 199.","DOI":"10.1093\/plphys\/kiaf540"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1016\/j.ccell.2015.12.004","article-title":"An integrated metabolic atlas of clear cell renal cell carcinoma","volume":"29","author":"Hakimi","year":"2016","journal-title":"Cancer Cell"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis of relative gene expression data using real-time quantitative PCR and the 2\u2212\u0394\u0394CT method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"}],"container-title":["Genes"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4425\/17\/5\/512\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T13:23:17Z","timestamp":1779196997000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4425\/17\/5\/512"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,26]]},"references-count":56,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2026,5]]}},"alternative-id":["genes17050512"],"URL":"https:\/\/doi.org\/10.3390\/genes17050512","relation":{},"ISSN":["2073-4425"],"issn-type":[{"value":"2073-4425","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,26]]}}}