{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T09:03:34Z","timestamp":1789981414363,"version":"4.0.1"},"reference-count":61,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,9,18]],"date-time":"2026-09-18T00:00:00Z","timestamp":1789689600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100012554","name":"Excellent Young and Middle-aged Scientific and Technological Innovation Team of Colleges and Universities in Hubei Province","doi-asserted-by":"publisher","award":["T2022050"],"award-info":[{"award-number":["T2022050"]}],"id":[{"id":"10.13039\/100012554","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32372227"],"award-info":[{"award-number":["32372227"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"award":["32372227"],"award-info":[{"award-number":["32372227"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Henan Provincial Science and Technology Key Research Project","award":["262102111103"],"award-info":[{"award-number":["262102111103"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Genes"],"abstract":"<jats:p>Background: The bHLH transcription factor family plays crucial roles in plant secondary metabolism, yet its genome-wide characterization and regulatory functions in flavonoid biosynthesis remain unexplored in Platycodon grandiflorus, a medicinal species with bioactive flavonoids. This study aims to systematically identify PgbHLH genes and screen for candidate regulators of flavonoid production. Methods: We performed genome-wide identification of bHLH family members using curated reference genomes, followed by phylogenetic classification, chromosomal localization, gene structure analysis, and collinearity assessment. Promoter cis-element prediction and subcellular localization assays for selected proteins were conducted. Expression profiles under methyl jasmonate (MeJA) treatment were evaluated via transcriptome sequencing and RT-qPCR. Protein\u2013protein interaction prediction, expression correlation, and weighted gene co-expression network analysis (WGCNA) were integrated to link PgbHLH genes with flavonoid structural genes. Results: A total of 95 PgbHLH genes were identified and clustered into 23 subfamilies, unevenly distributed across nine chromosomes. All members retained conserved bHLH domains but varied in exon\u2013intron structures. Thirty-six duplication events were detected, with Ka\/Ks ratios indicating purifying selection. Promoters harbored multiple hormone- and stress-responsive elements. PgbHLH37, PgbHLH42, and PgbHLH70 were confirmed to be nuclear-localized. Transcriptomic and RT-qPCR data showed differential expression of PgbHLH genes in response to MeJA. Integrative analyses identified 23 PgbHLH members associated with flavonoid biosynthetic genes, among which PgbHLH37, PgbHLH42, and PgbHLH70 exhibited strong multi-dimensional correlations and were nominated as high-priority candidate regulators. Conclusions: This systematic characterization provides a foundation for understanding bHLH-mediated flavonoid metabolism in P. grandiflorus, with three prioritized candidates for future functional validation. The findings are strictly supported by the presented genomic and expression evidence. However, our conclusions are based on total flavonoid content, and analysis of individual flavonoid species is required to determine pathway-specific regulation.<\/jats:p>","DOI":"10.3390\/genes17091144","type":"journal-article","created":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T08:03:47Z","timestamp":1789977827000},"page":"1144","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Genome-Wide Identification of bHLH Transcription Factors and Screening of Candidate Regulators Associated with Flavonoid Biosynthesis in Platycodon grandiflorus Under Methyl Jasmonate Treatment"],"prefix":"10.3390","volume":"17","author":[{"given":"Yalan","family":"Feng","sequence":"first","affiliation":[{"name":"College of Life Sciences, Wuchang College of Technology, Wuhan 430200, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"}]},{"given":"Runlong","family":"Wei","sequence":"additional","affiliation":[{"name":"College of Life Sciences, Wuchang College of Technology, Wuhan 430200, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"software","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"}]},{"given":"Yeying","family":"Wu","sequence":"additional","affiliation":[{"name":"College of Life Sciences, Wuchang College of Technology, Wuhan 430200, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"given":"Dandan","family":"Cheng","sequence":"additional","affiliation":[{"name":"College of Life Sciences, Wuchang College of Technology, Wuhan 430200, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"}]},{"given":"Zhao","family":"Zhao","sequence":"additional","affiliation":[{"name":"College of Life Sciences, Wuchang College of Technology, Wuhan 430200, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"software","vocabulary":"credit"}]},{"given":"Chao","family":"Ma","sequence":"additional","affiliation":[{"name":"College of Agriculture, Henan University of Science and Technology, Luoyang 471023, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,18]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"94","DOI":"10.1111\/j.1365-313X.2010.04459.x","article-title":"Evolutionary and comparative analysis of MYB and bHLH plant transcription factors","volume":"66","author":"Feller","year":"2011","journal-title":"Plant J."},{"key":"ref_2","first-page":"1333","article-title":"Role and functioning of bHLH transcription factors in jasmonate signalling","volume":"68","author":"Jonas","year":"2016","journal-title":"J. Exp. Bot."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Millard, P.S., Kragelund, B.B., and Burow, M. (2021). Evolution of A bHLH Interaction Motif. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22010447"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"226","DOI":"10.1186\/gb-2004-5-6-226","article-title":"An overview of the basic helix-loop-helix proteins","volume":"5","author":"Jones","year":"2004","journal-title":"Genome Biol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"7092","DOI":"10.1073\/pnas.86.18.7092","article-title":"Lc, a member of the maize R gene family responsible for tissue-specific anthocyanin production, encodes a protein similar to transcriptional activators and contains the myc-homology region","volume":"86","author":"Ludwig","year":"1989","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Zhang, T.T., Lv, W., Zhang, H.S., Ma, L., Li, P.H., Ge, L., and Li, G. (2018). Genome-wide analysis of the basic Helix-Loop-Helix (bHLH) transcription factor family in maize. BMC Plant Biol., 18.","DOI":"10.1186\/s12870-018-1441-z"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"735","DOI":"10.1093\/molbev\/msg088","article-title":"The basic helix-loop-helix transcription factor family in plants: A genome-wide study of protein structure and functional diversity","volume":"20","author":"Heim","year":"2003","journal-title":"Mol. Biol. Evol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1167","DOI":"10.1104\/pp.106.080580","article-title":"Genome-wide analysis of Basic\/Helix-Loop-Helix transcription factor family in Rice and Arabidopsis","volume":"141","author":"Li","year":"2006","journal-title":"Plant Physiol."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Guo, X.J., and Wang, J.R. (2017). Global identification, structural analysis and expression characterization of bHLH transcription factors in wheat. BMC Plant Biol., 17.","DOI":"10.1186\/s12870-017-1038-y"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.diff.2010.02.003","article-title":"Basic helix-loop-helix transcription factor gene family phylogenetics and nomenclature","volume":"80","author":"Skinner","year":"2010","journal-title":"Differentiation"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1038\/s41586-023-06282-3","article-title":"Al XYNE: Cooperation between bHLH transcription factors and histones for DNA access","volume":"619","author":"Michael","year":"2023","journal-title":"Nature"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"2465","DOI":"10.1093\/jxb\/erq442","article-title":"Recent advances in the transcriptional regulation of the flavonoid biosynthetic pathway","volume":"62","author":"Hichri","year":"2011","journal-title":"J. Exp. Bot."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1349","DOI":"10.1093\/genetics\/156.3.1349","article-title":"GL3 encodes a bHLH protein that regulates trichome development in Arabidopsis through interaction with GL1 and TTG1","volume":"156","author":"Payne","year":"2000","journal-title":"Genetics"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"4859","DOI":"10.1242\/dev.00681","article-title":"A network of redundant bHLH proteins functions in all TTG1-dependent pathways of Arabidopsis","volume":"130","author":"Zhang","year":"2003","journal-title":"Development"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"768","DOI":"10.1111\/j.1365-313X.2006.02733.x","article-title":"TT8 controls its own expression in a feedback regulation involving TTG1 and homologous MYB and bHLH factors, allowing a strong and cell-specific accumulation of flavonoids in Arabidopsis thaliana","volume":"46","author":"Baudry","year":"2006","journal-title":"Plant J."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Cao, Y.P., Li, K., Li, Y.L., Zhao, X.P., and Wang, L.H. (2020). MYB transcription factors as regulators of secondary metabolism in plants. Biology, 9.","DOI":"10.3390\/biology9030061"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"396","DOI":"10.1016\/j.plaphy.2020.06.015","article-title":"PsbHLH1, a novel transcription factor involved in regulating anthocyanin biosynthesis in tree peony (Paeonia suffruticosa)","volume":"154","author":"Qi","year":"2020","journal-title":"Plant Physiol. Biochem."},{"key":"ref_18","first-page":"1014","article-title":"Abnormal expression of bHLH3 disrupts a flavonoid homeostasis network, causing differences in pigment composition among mulberry fruits","volume":"7","author":"Li","year":"2023","journal-title":"Hortic. Res."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"891","DOI":"10.1007\/s00299-020-02537-9","article-title":"Expression of the subgroup IIIf bHLH transcription factor CpbHLH1 from Chimonanthus praecox (L.) in transgenic model plants inhibits anthocyanin accumulation","volume":"39","author":"Zhao","year":"2020","journal-title":"Plant Cell Rep."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1016\/j.bbrc.2014.05.139","article-title":"A novel bHLH transcription factor PebHLH35 from Populus euphratica confers drought tolerance through regulating stomatal development, photosynthesis and growth in Arabidopsis","volume":"450","author":"Dong","year":"2014","journal-title":"Biochem Biophys. Res. Commun."},{"key":"ref_21","first-page":"2801","article-title":"Pepper bHLH transcription factor CabHLH035 contributes to salt tolerance by modulating ion homeostasis and proline biosynthesis","volume":"9","author":"Zhang","year":"2023","journal-title":"Hortic. Res."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"224","DOI":"10.1016\/j.plaphy.2021.11.027","article-title":"Identification and function analysis of bHLH genes in response to cold stress in sweetpotato","volume":"169","author":"Jin","year":"2021","journal-title":"Plant Physiol. Biochem."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1102","DOI":"10.1111\/jipb.13248","article-title":"The jasmonate-induced bHLH gene SlJIG functions in terpene biosynthesis and resistance to insects and fungus","volume":"64","author":"Cao","year":"2022","journal-title":"J. Integr. Plant Biol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"uhae058","DOI":"10.1093\/hr\/uhae058","article-title":"RNA sequencing analysis reveals PgbHLH28 as the key regulator in response to methyl jasmonate-induced saponin accumulation in Platycodon grandifloras","volume":"11","author":"Zhang","year":"2024","journal-title":"Hortic. Res."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"154422","DOI":"10.1016\/j.phymed.2022.154422","article-title":"Platycodon grandiflorum (Jacq.) A. DC.: A review of phytochemistry, pharmacology, toxicology and traditional use","volume":"106","author":"Zhang","year":"2022","journal-title":"Phytomedicine"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"412","DOI":"10.1016\/j.fct.2018.11.001","article-title":"Platycodi Radix and its active compounds ameliorate against house dust mite-induced allergic airway inflammation and ER stress and ROS by enhancing anti-oxidation","volume":"123","author":"Lee","year":"2019","journal-title":"Food Chem. Toxicol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"11244","DOI":"10.1038\/srep11244","article-title":"Genome-wide characterisation and analysis of bHLH transcription factors related to tanshinone biosynthesis in Salvia miltiorrhiza","volume":"5","author":"Zhang","year":"2015","journal-title":"Sci. Rep."},{"key":"ref_28","first-page":"33","article-title":"TBtools-Domestic biological software in the era of big data","volume":"17","author":"Chen","year":"2022","journal-title":"Sci. Obs."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"e49","DOI":"10.1093\/nar\/gkr1293","article-title":"MCScanX: A toolkit for detection and evolutionary analysis of gene synteny and collinearity","volume":"40","author":"Wang","year":"2012","journal-title":"Nucleic Acids Res."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1194","DOI":"10.1016\/j.molp.2020.06.009","article-title":"TBtools: An integrative toolkit developed for interactive analyses of big biological data","volume":"13","author":"Chen","year":"2020","journal-title":"Mol. Plant"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1038\/sj.cdd.4401712","article-title":"AtBAG6, a novel calmodulin-binding protein, induces programmed cell death in yeast and plants","volume":"13","author":"Kang","year":"2006","journal-title":"Cell Death Differ."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"156","DOI":"10.1038\/s41438-021-00590-3","article-title":"The R2R3-type MYB transcription factor MdMYB90-like is responsible for the enhanced skin color of an apple bud sport mutant","volume":"8","author":"Sun","year":"2021","journal-title":"Hortic. Res."},{"key":"ref_33","first-page":"963","article-title":"Cloning and Analysis of a Basic Helix-loop-helix (bHLH) Transcription Factor MtbHLH148 from Medicago truncatula L.","volume":"39","author":"Juping","year":"2019","journal-title":"Acta Bot. Boreali-Occident. Sin."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1159","DOI":"10.1007\/s11240-014-0671-5","article-title":"Regeneration of buckwheat plantlets from hypocotyl and the influence of exogenous hormones on rutin content and rutin biosynthetic gene expression in vitro","volume":"120","author":"Hou","year":"2015","journal-title":"Plant Cell Tiss. Org."},{"key":"ref_35","first-page":"1040","article-title":"Effects of Methyl Jasmonate Accumulation of Flavonoids and Related Gene Expression of Buckwheat Sprouts","volume":"34","author":"Luo","year":"2015","journal-title":"Genom. Appl. Biol."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Deng, X.J., Zheng, F.X., Xu, Z.T., Mao, X.P., Yu, Z.M., and Shen, X.X. (2025). Comprehensive Identification and Abscisic Acid-Responsive Expression Profiling of NAC Transcription Factor in Triterpenoid Saponin in Hedera helix. Biomolecules, 15.","DOI":"10.3390\/biom15111557"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Xiang, L., He, P., Shu, G.P., Yuan, M.Y., Wen, M.L., Lan, X.Z., Liao, Z.H., and Tang, Y.L. (2022). AabHLH112, a bHLH transcription factor, positively regulates sesquiterpenes biosynthesis in Artemisia annua. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.973591"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Ullah, I., Yuan, W.B., Uzair, M., Li, S.S., Rehman, O.U., Nanda, S., and Wu, H.Z. (2022). Molecular characterization of bHLH transcription factor family in Rose (Rosa chinensis Jacq.) under Botrytis cinerea infection. Horticulturae, 8.","DOI":"10.3390\/horticulturae8110989"},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Hao, Y.Q., Zong, X.M., Ren, P., Qian, Y.Q., and Fu, A.G. (2021). Basic Helix-Loop-Helix (bHLH) transcription factors regulate a wide range of functions in Arabidopsis. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22137152"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"37636","DOI":"10.1074\/jbc.M605895200","article-title":"A R2R3 type MYB transcription factor is involved in the cold regulation of CBF genes and in acquired freezing tolerance","volume":"281","author":"Agarwal","year":"2006","journal-title":"J. Biol. Chem."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"592","DOI":"10.1111\/j.1365-313X.2006.02980.x","article-title":"ABA induction of miR159 controls transcript levels of two MYB factors during Arabidopsis seed germination","volume":"49","author":"Reyes","year":"2007","journal-title":"Plant J."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Wang, Y., and Liu, A.Z. (2020). Genomic characterization and expression analysis of Basic Helix-Loop-Helix (bHLH) family genes in traditional chinese herb Dendrobium officinale. Plants, 9.","DOI":"10.3390\/plants9081044"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Yang, J.H., Zhang, B., Gu, G., Yuan, J.Z., Shen, S.J., Jin, L., Lin, Z.Q., Lin, J.F., and Xie, X.F. (2022). Genome-wide identification and expression analysis of the R2R3-MYB gene family in tobacco (Nicotiana tabacum L.). BMC Genom., 23.","DOI":"10.1186\/s12864-022-08658-7"},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Dossa, K., Diouf, D., and Ciss\u00e9, N. (2016). Genome-wide investigation of Hsf genes in sesame reveals their segmental duplication expansion and their active role in drought stress response. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.01522"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1398","DOI":"10.1104\/pp.110.153593","article-title":"Genome-wide classification and evolutionary analysis of the bHLH family of transcription factors in Arabidopsis, Poplar, Rice, Moss, and Algae1","volume":"153","author":"Galstyan","year":"2010","journal-title":"Plant Physiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1016\/S1360-1385(00)01600-9","article-title":"The WRKY superfamily of plant transcription factors","volume":"5","author":"Eulgem","year":"2000","journal-title":"Trends Plant Sci."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Hou, L.X., Yuan, X.C., Li, Z.H., and Li, J.R. (2021). Genome-wide identification of CYP72A gene family and expression patterns related to jasmonic acid treatment and steroidal saponin accumulation in Dioscorea zingiberensis. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms222010953"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1111\/j.1365-313X.2011.04575.x","article-title":"The basic helix-loop-helix transcription factor CrMYC2 controls the jasmonate-responsive expression of the ORCA genes that regulate alkaloid biosynthesis in Catharanthus roseus","volume":"67","author":"Zhang","year":"2011","journal-title":"Plant J."},{"key":"ref_49","first-page":"193","article-title":"A bHLH gene from Tamarix hispida improves abiotic stress tolerance by enhancing osmotic potential and decreasing reactive oxygen species accumulation","volume":"36","author":"Ji","year":"2016","journal-title":"Tree Physiol."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Yue, M.L., Jiang, L.Y., Zhang, N.T., Zhang, L.X., Liu, Y.Q., Lin, Y.X., Zhang, Y.T., Luo, Y., Zhang, Y., and Wang, Y. (2023). Regulation of flavonoids in strawberry fruits by FaMYB5\/FaMYB10 dominated MYB-bHLH-WD40 ternary complexes. Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1145670"},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Wang, M.J., Ou, Y., Li, Z., Zheng, Q.D., Ke, Y.J., Lai, H.P., Lan, S.R., Peng, D.H., Liu, Z.J., and Ai, Y. (2023). Genome-wide identification and analysis of bHLH transcription factors related to anthocyanin biosynthesis in Cymbidium ensifolium. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24043825"},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Yin, W., Wang, L., Shu, Q.Q., Chen, M.J., Li, F., and Luo, X.B. (2024). Genome-wide identification and expression analysis of the CONSTANS-like family in potato (Solanum tuberosum L.). Front. Genet., 15.","DOI":"10.3389\/fgene.2024.1390411"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1016\/j.gene.2019.02.063","article-title":"Revealing shared differential co-expression profiles in rice infected by virus from reoviridae and sequiviridae group","volume":"25","author":"Sahu","year":"2019","journal-title":"Gene"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1016\/j.plaphy.2019.07.023","article-title":"Comparative transcriptomics and weighted gene co-expression correlation network analysis (WGCNA) reveal potential regulation mechanism of carotenoid accumulation in Chrysanthemum \u00d7 morifolium","volume":"142","author":"Lu","year":"2019","journal-title":"Plant Physiol. Biochem."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1007\/s11103-019-00945-4","article-title":"Transcriptome and weighted correlation network analyses provide insights into inflorescence stem straightness in Paeonia lactiflora","volume":"102","author":"Wan","year":"2020","journal-title":"Plant Mol. Biol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1093\/hr\/uhac198","article-title":"A bHLH transcription factor, SlbHLH96, promotes drought tolerance in tomato","volume":"9","author":"Liang","year":"2022","journal-title":"Hortic. Res."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"906","DOI":"10.1093\/oxfordjournals.pcp.a029038","article-title":"Analysis of the promoter of the auxin-inducible gene, parC, of tobacco","volume":"37","author":"Sakai","year":"1996","journal-title":"Plant Cell Physiol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"13723","DOI":"10.1038\/s41598-020-69305-3","article-title":"Genome-wide identification and characterization of bHLH family genes from Ginkgo biloba","volume":"10","author":"Zhou","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_59","first-page":"13","article-title":"Basic helix-loop-helix(bHLH) transcription factor MdbHLH3 negatively affects the storage performance of postharvest apple fruit","volume":"8","author":"Wang","year":"2022","journal-title":"Hortic. Plant J."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1046\/j.1365-313X.2003.01708.x","article-title":"Interaction between two cis-acting elements, ABRE and DRE, in ABA-dependent expression of Arabidopsis rd29A gene in response to dehydration and high-salinity stresses","volume":"34","author":"Narusaka","year":"2003","journal-title":"Plant J."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1461","DOI":"10.1016\/j.molp.2017.08.007","article-title":"The bHLH transcription factors MYC2, MYC3, and MYC4 are required for jasmonate-mediated inhibition of flowering in Arabidopsis","volume":"10","author":"Wang","year":"2017","journal-title":"Mol. 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