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Therefore, identifying novel PCZ targets and breeding PCZ-free cultivars is critical for the safe production of flowering Chinese cabbage. Here, we identified three P4-ATPase flippase homologues aminophospholipid ATPase 3 (BraALA3a\/b\/c) in flowering Chinese cabbage that function as sensitive targets for PCZ. These proteins exhibit high binding affinity for PCZ, which directly inhibits their ATPase activity. Overexpression of the BraALA3 homologues enhanced plant growth and increased sensitivity to PCZ, whereas knockdown led to dwarfism and reduced sensitivity. Based on these findings, we identified editable active sites via protoplast-based screening. Genetic transformation of one such site yielded BraALA3a\/braala3aK200T mutant lines, which displayed a dwarf and compact architecture. These findings provide a precise molecular target for developing PCZ-free germplasm in flowering Chinese cabbage through gene editing.<\/jats:p>","DOI":"10.1021\/acs.jafc.6c02868","type":"journal-article","created":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T20:02:05Z","timestamp":1786651325000},"page":"25935-25949","source":"Crossref","is-referenced-by-count":0,"title":["BraALA3 Homologues Mediate Propiconazole Effects on\nGrowth of Flowering Chinese Cabbage"],"prefix":"10.1021","volume":"74","author":[{"given":"Qing","family":"Gao","sequence":"first","affiliation":[{"name":"South China Agricultural University , , ,","place":["Guangzhou, China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yunxue","family":"Song","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , ,","place":["Guangzhou, China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yongsheng","family":"Yang","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , ,","place":["Guangzhou, China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Siwei","family":"Wang","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , ,","place":["Guangzhou, China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiancong","family":"Ruan","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , ,","place":["Guangzhou, China, 510642"]},{"name":"Guangzhou Shennong Modern Agriculture Co., Ltd. , ,","place":["Guangzhou, China, 511453"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhicheng","family":"Liu","sequence":"additional","affiliation":[{"name":"Guangzhou Shennong Modern Agriculture Co., Ltd. , ,","place":["Guangzhou, China, 511453"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dekang","family":"Guo","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , ,","place":["Guangzhou, China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yiping","family":"Chen","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , ,","place":["Guangzhou, China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinping","family":"Wang","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , ,","place":["Guangzhou, China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ronghua","family":"Chen","sequence":"additional","affiliation":[{"name":"Guangdong Academy of Sciences , , ,","place":["Guangzhou, China, 510316"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7841-2396","authenticated-orcid":true,"given":"Hanhong","family":"Xu","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , ,","place":["Guangzhou, China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2321-889X","authenticated-orcid":true,"given":"Fei","family":"Lin","sequence":"additional","affiliation":[{"name":"South China Agricultural University , , ,","place":["Guangzhou, China, 510642"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,8,13]]},"reference":[{"key":"2026090104472961700_cit1","doi-asserted-by":"publisher","first-page":"129535","DOI":"10.1016\/j.foodchem.2021.129535","article-title":"Nutritional\nmetabolites in Brassica rapa subsp. chinensis var. parachinensis (choy sum) at three different\ngrowth stages: Microgreen, seedling and adult plant","volume":"357","author":"Zou","year":"2021","journal-title":"Food Chem."},{"issue":"18","key":"2026090104472961700_cit2","doi-asserted-by":"publisher","first-page":"5318","DOI":"10.1021\/acs.jafc.1c00155","article-title":"Light Intensity Plays\nContrasting Roles in Regulating Metabolite Compositions in Choy Sum\n(Brassica rapa var. parachinensis)","volume":"69","author":"Huang","year":"2021","journal-title":"J. Agric. Food Chem."},{"issue":"13","key":"2026090104472961700_cit3","doi-asserted-by":"publisher","first-page":"7412","DOI":"10.3390\/ijms23137412","article-title":"Identification of BcARR Genes and\nCTK Effects on Stalk Development of Flowering Chinese Cabbage","volume":"23","author":"Ou","year":"2022","journal-title":"Int. J. Mol. Sci."},{"issue":"2","key":"2026090104472961700_cit4","doi-asserted-by":"publisher","first-page":"uhae299","DOI":"10.1093\/hr\/uhae299","article-title":"Genomic selection and genetic architecture of agronomic traits during\nmodern flowering Chinese cabbage breeding","volume":"12","author":"Zhao","year":"2025","journal-title":"Hortic.\nRes."},{"issue":"8","key":"2026090104472961700_cit5","doi-asserted-by":"publisher","first-page":"uhad119","DOI":"10.1093\/hr\/uhad119","article-title":"Role of BraRGL1 in regulation of Brassica rapa bolting and flowering","volume":"10","author":"Wang","year":"2023","journal-title":"Hortic.\nRes."},{"issue":"1","key":"2026090104472961700_cit6","doi-asserted-by":"publisher","first-page":"5470","DOI":"10.1038\/s41467-024-49721-z","article-title":"Temperature-dependent\njumonji demethylase modulates\nflowering time by targeting H3K36me2\/3 in Brassica rapa","volume":"15","author":"Xin","year":"2024","journal-title":"Nat. Commun."},{"issue":"3","key":"2026090104472961700_cit7","first-page":"47","article-title":"Studies on\nEffect of Propiconazole on Plant Height and Yield of Flowering Chinese\nCabbage and Its Residue Behavior","volume":"1","author":"Huang","year":"2019","journal-title":"China Veg."},{"issue":"4","key":"2026090104472961700_cit8","first-page":"752","article-title":"Residue and Dietary Exposure Risk\nAssessment of Propiconazole and\nChlormequat in ChineseKale, Flowering Chinese Cabbage and Pak-Choi\nin Guangzhou","volume":"38","author":"Ye","year":"2017","journal-title":"Chin. J. Top. Crops"},{"issue":"50","key":"2026090104472961700_cit9","doi-asserted-by":"publisher","first-page":"27775","DOI":"10.1021\/acs.jafc.4c06923","article-title":"Stereoselective In\nVitro Metabolism, Hepatotoxicity, and Cytotoxic\nEffects of Four Enantiomers of the Fungicide Propiconazole","volume":"72","author":"Ma","year":"2024","journal-title":"J. Agric. Food Chem."},{"issue":"26","key":"2026090104472961700_cit10","doi-asserted-by":"publisher","first-page":"7399","DOI":"10.1021\/acs.jafc.1c01086","article-title":"In Vitro and In Vivo Study on the\nToxic Effects of Propiconazole Fungicide in the Pathogenesis of Liver\nFibrosis","volume":"69","author":"Kwon","year":"2021","journal-title":"J. Agric. Food Chem."},{"issue":"2","key":"2026090104472961700_cit11","doi-asserted-by":"publisher","first-page":"359","DOI":"10.1128\/AEM.02159-07","article-title":"The Cytochrome P450\nLanosterol 14\u03b1-Demethylase\nGene Is a Demethylation Inhibitor Fungicide Resistance Determinant\nin Monilinia fructicola Field Isolates from Georgia","volume":"74","author":"Luo","year":"2008","journal-title":"Appl. Environ. Microbiol."},{"issue":"3","key":"2026090104472961700_cit12","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1289\/ehp.5785","article-title":"Azole fungicides\naffect mammalian steroidogenesis by inhibiting sterol 14 alpha-demethylase\nand aromatase","volume":"111","author":"Zarn","year":"2003","journal-title":"Environ. Health Perspect."},{"issue":"2","key":"2026090104472961700_cit13","doi-asserted-by":"publisher","first-page":"734","DOI":"10.1104\/pp.105.066639","article-title":"Lipid Signaling\nin Plants. Cloning and Expression Analysis of the Obtusifoliol 14\u03b1-Demethylase\nfrom Solanum chacoense Bitt., a Pollination- and\nFertilization-Induced Gene with Both Obtusifoliol and Lanosterol Demethylase\nActivity","volume":"139","author":"O\u2019Brien","year":"2005","journal-title":"Plant Physiol."},{"issue":"5","key":"2026090104472961700_cit14","doi-asserted-by":"publisher","DOI":"10.1080\/15592324.2016.1158372","article-title":"In vitro and\nin vivo evidence for the inhibition of brassinosteroid synthesis by\npropiconazole through interference with side chain hydroxylation","volume":"11","author":"Oh","year":"2016","journal-title":"Plant Signaling Behav."},{"issue":"4","key":"2026090104472961700_cit15","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1007\/bf00228939","article-title":"The effects of a triazole fungicide,\npropiconazole, on pollen germination, tube growth and cytoskeletal\ndistribution in Tradescantia virginiana","volume":"8","author":"He","year":"1995","journal-title":"Sex. Plant Reprod."},{"key":"2026090104472961700_cit16","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1016\/j.plipres.2016.08.003","article-title":"Lipid somersaults: Uncovering the mechanisms of protein-mediated\nlipid flipping","volume":"64","author":"Pomorski","year":"2016","journal-title":"Prog. Lipid Res."},{"issue":"8","key":"2026090104472961700_cit17","doi-asserted-by":"publisher","first-page":"1068","DOI":"10.1016\/j.bbalip.2011.12.007","article-title":"Phospholipid\nflippases:\nBuilding asymmetric membranes and transport vesicles","volume":"1821","author":"Sebastian","year":"2012","journal-title":"Biochim. Biophys. Acta Gen. 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Agric. Sci."},{"issue":"10","key":"2026090104472961700_cit28","doi-asserted-by":"publisher","first-page":"1888","DOI":"10.1111\/pbi.13869","article-title":"An amino acid transporter-like protein\n(OsATL15) facilitates the systematic distribution of thiamethoxam\nin rice for controlling the brown planthopper","volume":"20","author":"Xiao","year":"2022","journal-title":"Plant Biotechnol. J."},{"issue":"7","key":"2026090104472961700_cit29","doi-asserted-by":"publisher","first-page":"1870","DOI":"10.1093\/molbev\/msw054","article-title":"MEGA7: Molecular Evolutionary Genetics\nAnalysis Version 7.0 for Bigger Datasets","volume":"33","author":"Kumar","year":"2016","journal-title":"Mol.\nBiol. Evol."},{"issue":"5","key":"2026090104472961700_cit30","doi-asserted-by":"publisher","first-page":"1218","DOI":"10.1111\/tpj.16418","article-title":"The herbicide bensulfuron-methyl\ninhibits rice seedling development by blocking calcium ion flux in\nthe OsCNGC12 channel","volume":"116","author":"Guo","year":"2023","journal-title":"Plant J."},{"key":"2026090104472961700_cit31","doi-asserted-by":"publisher","first-page":"4347","DOI":"10.1038\/s41467-025-59582-9","article-title":"Fabricating\nsupramolecular pre-emergence herbicide CPAM-BPyHs for\nfarming herbicide-resistant rice","volume":"16","author":"Chen","year":"2025","journal-title":"Nat. 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Mol.\nSci."},{"issue":"2","key":"2026090104472961700_cit38","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1016\/j.softx.2015.06.001","article-title":"GROMACS: High\nperformance\nmolecular simulations through multi-level parallelism from laptops\nto supercomputers","volume":"1","author":"Abraham","year":"2015","journal-title":"SoftwareX"},{"key":"2026090104472961700_cit39","doi-asserted-by":"publisher","first-page":"146505","DOI":"10.1016\/j.foodchem.2025.146505","article-title":"Identification of new coumarin-cinnamic\nacid hybrid FtsZ inhibitors as potential bactericides for enhancing\nthe yield and quality of rice","volume":"495","author":"Zhang","year":"2025","journal-title":"Food Chem."},{"issue":"6","key":"2026090104472961700_cit40","doi-asserted-by":"publisher","first-page":"458","DOI":"10.1007\/s13238-020-00712-y","article-title":"Structures of a P4-ATPase lipid flippase\nin lipid bilayers","volume":"11","author":"He","year":"2020","journal-title":"Protein Cell"},{"key":"2026090104472961700_cit41","doi-asserted-by":"publisher","first-page":"113242","DOI":"10.1016\/j.ejmech.2021.113242","article-title":"Discovery and structural\noptimization of 3-O-\u03b2-chacotriosyl\noleanane-type triterpenoids as potent entry inhibitors of SARS-CoV-2\nvirus infections","volume":"215","author":"Li","year":"2021","journal-title":"Eur. J. Med. Chem."},{"key":"2026090104472961700_cit42","doi-asserted-by":"publisher","first-page":"142770","DOI":"10.1016\/j.foodchem.2025.142770","article-title":"Sodium hydrosulfide\napplication induces chilling tolerance in banana fruits by enhancing\nantioxidant gene expression through the upregulation of the ethylene\nresponse factors MaERF53L\/121L","volume":"471","author":"Lin","year":"2025","journal-title":"Food Chem."},{"issue":"10","key":"2026090104472961700_cit43","doi-asserted-by":"publisher","first-page":"3424","DOI":"10.3390\/ijms21103424","article-title":"Complex Molecular Evolution\nand Expression of Expansin Gene Families\nin Three Basic Diploid Species of Brassica","volume":"21","author":"Liu","year":"2020","journal-title":"Int. J. Mol. Sci."},{"issue":"5","key":"2026090104472961700_cit44","doi-asserted-by":"publisher","first-page":"752","DOI":"10.1089\/crispr.2021.0045","article-title":"Efficient and Economical\nTargeted Insertion in Plant Genomes via Protoplast Regeneration","volume":"4","author":"Hsu","year":"2021","journal-title":"CRISPR J."},{"issue":"7","key":"2026090104472961700_cit45","doi-asserted-by":"publisher","first-page":"1295","DOI":"10.1111\/pbi.12870","article-title":"Application\nof protoplast\ntechnology to CRISPR\/Cas9 mutagenesis: from single-cell mutation detection\nto mutant plant regeneration","volume":"16","author":"Lin","year":"2018","journal-title":"Plant Biotechnol.\nJ."},{"issue":"4","key":"2026090104472961700_cit46","doi-asserted-by":"publisher","first-page":"690","DOI":"10.1016\/j.molp.2025.03.002","article-title":"SuperDecode: An integrated toolkit\nfor analyzing mutations induced by genome editing","volume":"18","author":"Li","year":"2025","journal-title":"Mol. 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Res."},{"issue":"1","key":"2026090104472961700_cit48","doi-asserted-by":"publisher","first-page":"15517","DOI":"10.1038\/s41598-017-15699-6","article-title":"Transcriptomic\nanalysis of the regulation\nof stalk development in flowering Chinese cabbage (Brassica campestris) by RNA sequencing","volume":"7","author":"Huang","year":"2017","journal-title":"Sci.\nRep."},{"issue":"19","key":"2026090104472961700_cit49","doi-asserted-by":"publisher","first-page":"7954","DOI":"10.1074\/jbc.M116.751487","article-title":"High phosphatidylinositol 4-phosphate\n(PI4P)-dependent\nATPase activity for the Drs2p-Cdc50p flippase after removal of its\nN- and C-terminal extensions","volume":"292","author":"Azouaoui","year":"2017","journal-title":"J. Biol. 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