{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T20:06:58Z","timestamp":1792786018270,"version":"4.5.0"},"reference-count":78,"publisher":"Oxford University Press (OUP)","issue":"17","license":[{"start":{"date-parts":[[2026,6,17]],"date-time":"2026-06-17T00:00:00Z","timestamp":1781654400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2022YFC3501700"],"award-info":[{"award-number":["2022YFC3501700"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2023YFC3504800"],"award-info":[{"award-number":["2023YFC3504800"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["U23A20512"],"award-info":[{"award-number":["U23A20512"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32170402"],"award-info":[{"award-number":["32170402"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012247","name":"Program of Shanghai Academic\/Technology Research Leader","doi-asserted-by":"publisher","award":["23XD1423500"],"award-info":[{"award-number":["23XD1423500"]}],"id":[{"id":"10.13039\/501100012247","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003399","name":"Science and Technology Commission of Shanghai Municipality","doi-asserted-by":"publisher","award":["24DZ2306100"],"award-info":[{"award-number":["24DZ2306100"]}],"id":[{"id":"10.13039\/501100003399","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Organizational Key Research and Development Program of Shanghai University of Traditional Chinese Medicine","award":["2023YZZ01"],"award-info":[{"award-number":["2023YZZ01"]}],"id":[{"id":"https:\/\/ror.org\/00z27jk27","id-type":"ROR","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100004001","name":"Guizhou Science and Technology Department","doi-asserted-by":"publisher","award":["CXPTXM[2025]010\/[2025]021\/KXJZ[2025]014"],"award-info":[{"award-number":["CXPTXM[2025]010\/[2025]021\/KXJZ[2025]014"]}],"id":[{"id":"10.13039\/501100004001","id-type":"DOI","asserted-by":"publisher"}]},{"name":"The Shanghai Committee of Science and Technology","award":["25HC2810300"],"award-info":[{"award-number":["25HC2810300"]}]},{"name":"Chinese medicine resources","award":["2060302"],"award-info":[{"award-number":["2060302"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,16]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Sweet wormwood (Artemisia annua) produces and deposits artemisinin in its glandular trichomes but the molecular mechanisms of trichome development remain poorly understood. Here, we conduct single-nucleus RNA sequencing of 10-day-old A. annua seedlings, revealing the specific RNA expression of the leaf trichome cells. The reconstruction of the developmental trajectory of trichome cells not only identifies novel expressed genes, but also elucidates trichome development involved in photosynthesis, auxin biosynthesis, and cutin biosynthesis. By integrating the developmental trajectory of trichome cells with histochemical assays, we identified several genes not previously reported to be involved in trichome development, such as TOE3, MYB1, WRKY10, and ZNF. Moreover, we identified a gene in subcluster 0 encoding a previously unknown WD40 repeat nuclear protein, WDR1, potentially involved in trichome development. We showed that WDR1 not only interacts with the transcription factor SPL9 and enhances its ability to activate HD1 expression, but also directly binds to the transcription factor HD1 itself, thereby forming a regulatory feedback loop that modulates trichome development in A. annua. This study reveals a previously uncharacterized gene that regulates trichome development in A. annua based on single-cell transcriptome analysis. The results presented here offer unprecedented insight into a new pathway for enhancing trichome density and artemisinin production.<\/jats:p>","DOI":"10.1093\/jxb\/erag287","type":"journal-article","created":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T11:53:49Z","timestamp":1781178829000},"page":"5715-5733","source":"Crossref","is-referenced-by-count":0,"title":["Single-nucleus RNA sequencing reveals the developmental trajectory and transcriptional regulatory networks of trichomes in\n                    <i>Artemisia annua<\/i>"],"prefix":"10.1093","volume":"77","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1965-8809","authenticated-orcid":false,"given":"Zongyou","family":"Lv","sequence":"first","affiliation":[{"name":"Research and Development Center of Chinese Medicine Resources and Biotechnology, Shanghai University of Traditional Chinese Medicine , Shanghai 201203 ,","place":["China"]},{"name":"State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs , Beijing 100700 ,","place":["China"]},{"name":"State Key Laboratory of Discovery and Utilization of Functional Components in Traditional Chinese Medicine, Institute of Chinese Materia Medica, Shanghai University of Traditional Chinese Medicine , Shanghai 201203 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chuhan","family":"Li","sequence":"additional","affiliation":[{"name":"Research and Development Center of Chinese Medicine Resources and Biotechnology, Shanghai University of Traditional Chinese Medicine , Shanghai 201203 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rui","family":"Jiang","sequence":"additional","affiliation":[{"name":"Research and Development Center of Chinese Medicine Resources and Biotechnology, Shanghai University of Traditional Chinese Medicine , Shanghai 201203 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2995-306X","authenticated-orcid":false,"given":"Xinlei","family":"Jia","sequence":"additional","affiliation":[{"name":"The Industrial Crop Institute, Shanxi Agricultural University\/Shanxi Academy of Agricultural Sciences , Shanxi 030031 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8537-9245","authenticated-orcid":false,"given":"Peng","family":"Di","sequence":"additional","affiliation":[{"name":"State Local Joint Engineering Research Center of Ginseng Breeding and Application, College of Chinese Medicinal Materials, Jilin Agricultural University , Jillin 130118 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0907-0190","authenticated-orcid":false,"given":"Ying","family":"Xiao","sequence":"additional","affiliation":[{"name":"Research and Development Center of Chinese Medicine Resources and Biotechnology, Shanghai University of Traditional Chinese Medicine , Shanghai 201203 ,","place":["China"]},{"name":"State Key Laboratory of Discovery and Utilization of Functional Components in Traditional Chinese Medicine, Institute of Chinese Materia Medica, Shanghai University of Traditional Chinese Medicine , Shanghai 201203 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0025-1315","authenticated-orcid":false,"given":"Wansheng","family":"Chen","sequence":"additional","affiliation":[{"name":"Research and Development Center of Chinese Medicine Resources and Biotechnology, Shanghai University of Traditional Chinese Medicine , Shanghai 201203 ,","place":["China"]},{"name":"State Key Laboratory of Discovery and Utilization of Functional Components in Traditional Chinese Medicine, Institute of Chinese Materia Medica, Shanghai University of Traditional Chinese Medicine , Shanghai 201203 ,","place":["China"]},{"name":"Department of Pharmacy, Changzheng Hospital, Second Military Medical University , Shanghai 200003 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,6,17]]},"reference":[{"key":"2026091519522645500_erag287-B1","doi-asserted-by":"crossref","first-page":"2036","DOI":"10.1111\/nph.17827","article-title":"Subtle interplay between trichome development and cuticle formation in plants","volume":"233","author":"Berhin","year":"2022","journal-title":"New Phytologist"},{"key":"2026091519522645500_erag287-B2","doi-asserted-by":"crossref","first-page":"1853","DOI":"10.1016\/j.cub.2010.09.046","article-title":"Keeping cell identity in Arabidopsis requires PRC1 RING-finger homologs that catalyze H2A monoubiquitination","volume":"20","author":"Bratzel","year":"2010","journal-title":"Current Biology"},{"key":"2026091519522645500_erag287-B3","doi-asserted-by":"crossref","first-page":"2303","DOI":"10.1038\/s41467-024-46683-0","article-title":"Spatiotemporal formation of glands in plants is modulated by MYB-like transcription factors","volume":"15","author":"Chang","year":"2024","journal-title":"Nature Communication"},{"key":"2026091519522645500_erag287-B4","first-page":"1156","article-title":"The WD-repeat protein CsTTG1 regulates fruit wart formation through interaction with the homeodomain-leucine zipper I protein Mict","volume":"171","author":"Chen","year":"2016","journal-title":"Plant Physiology"},{"key":"2026091519522645500_erag287-B5","doi-asserted-by":"crossref","first-page":"855","DOI":"10.1016\/j.molp.2021.05.002","article-title":"PlantscRNAdb: a database for plant single-cell RNA analysis","volume":"14","author":"Chen","year":"2021","journal-title":"Molecular Plant"},{"key":"2026091519522645500_erag287-B6","doi-asserted-by":"crossref","first-page":"304","DOI":"10.1111\/nph.14373","article-title":"GLANDULAR TRICHOME-SPECIFIC WRKY 1 promotes artemisinin biosynthesis in Artemisia annua","volume":"214","author":"Chen","year":"2017","journal-title":"New Phytologist"},{"key":"2026091519522645500_erag287-B7","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1038\/s41438-021-00652-6","article-title":"AaWRKY17, a positive regulator of artemisinin biosynthesis, is involved in resistance to Pseudomonas syringae in Artemisia annua","volume":"8","author":"Chen","year":"2021","journal-title":"Horticulture Research"},{"key":"2026091519522645500_erag287-B8","doi-asserted-by":"crossref","DOI":"10.1016\/j.indcrop.2025.120989","article-title":"IPPI, the core enzyme in C5-unit metabolism, regulates terpenoid biosynthesis in Artemisia annua","volume":"229","author":"Deng","year":"2025","journal-title":"Industrial Crops and Products"},{"key":"2026091519522645500_erag287-B9","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1046\/j.1365-313X.1996.10030393.x","article-title":"The Arabidopsis Athb-10 (GLABRA2) is an HD-Zip protein required for regulation of root hair development","volume":"10","author":"Di Cristina","year":"1996","journal-title":"The Plant Journal"},{"key":"2026091519522645500_erag287-B10","doi-asserted-by":"crossref","DOI":"10.1016\/j.xplc.2023.100554","article-title":"Single-cell RNA landscape of the special fiber initiation process in Bombax ceiba","volume":"4","author":"Ding","year":"2023","journal-title":"Plant Communications"},{"key":"2026091519522645500_erag287-B11","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1186\/s43897-024-00085-4","article-title":"TrichomeLess regulator 3 is required for trichome initial and cuticle biosynthesis in Artemisia annua","volume":"4","author":"Dong","year":"2024","journal-title":"Molecular Horticulture"},{"key":"2026091519522645500_erag287-B12","doi-asserted-by":"crossref","first-page":"6297","DOI":"10.1093\/jxb\/eraa344","article-title":"The HD-ZIP IV transcription factor Tril regulates fruit spine density through gene dosage effects in cucumber","volume":"71","author":"Du","year":"2020","journal-title":"Journal of Experimental Botany"},{"key":"2026091519522645500_erag287-B13","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1086\/297096","article-title":"Development and fine structure of the glandular trichomes of Artemisia annua L","volume":"154","author":"Duke","year":"1993","journal-title":"International Journal of Plant Sciences"},{"key":"2026091519522645500_erag287-B14","doi-asserted-by":"crossref","first-page":"1413","DOI":"10.1007\/s11434-016-1108-9","article-title":"Manipulation of SlMXl for enhanced carotenoids accumulation and drought resistance in tomato","volume":"61","author":"Ewas","year":"2016","journal-title":"Science Bulletin"},{"key":"2026091519522645500_erag287-B15","doi-asserted-by":"crossref","first-page":"1858","DOI":"10.1111\/nph.17453","article-title":"AaWRKY9 contributes to light- and jasmonate-mediated to regulate the biosynthesis of artemisinin in Artemisia annua","volume":"231","author":"Fu","year":"2021","journal-title":"New Phytologist"},{"key":"2026091519522645500_erag287-B16","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1038\/nprot.2007.13","article-title":"High-efficiency yeast transformation using the LiAc\/SS carrier DNA\/PEG method","volume":"2","author":"Gietz","year":"2007","journal-title":"Nature Protocols"},{"key":"2026091519522645500_erag287-B17","doi-asserted-by":"crossref","DOI":"10.1016\/j.plantsci.2021.111172","article-title":"AaSPL9 affects glandular trichomes initiation by positively regulating expression of AaHD1 in Artemisia annua L","volume":"317","author":"He","year":"2022","journal-title":"Plant Science"},{"key":"2026091519522645500_erag287-B18","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1007\/s11103-004-6768-1","article-title":"Two WD-repeat genes from cotton are functional homologues of the Arabidopsis thaliana TRANSPARENT TESTA GLABRA1 (TTG1) gene","volume":"57","author":"Humphries","year":"2005","journal-title":"Plant Molecular Biology"},{"key":"2026091519522645500_erag287-B19","doi-asserted-by":"crossref","DOI":"10.1016\/j.ijbiomac.2024.138834","article-title":"Genome-wide identification of the WD40 protein family and functional characterization of AaTTG1 in Artemisia annua","volume":"289","author":"Jiang","year":"2025","journal-title":"International Journal of Biological Macromolecules"},{"key":"2026091519522645500_erag287-B20","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1007\/s11105-013-0603-2","article-title":"Molecular cloning and characterization of a trichome-specific promoter of artemisinic aldehyde \u039411(13) reductase (DBR2) in Artemisia annua","volume":"32","author":"Jiang","year":"2014","journal-title":"Plant Molecular Biology Reporter"},{"key":"2026091519522645500_erag287-B21","doi-asserted-by":"crossref","first-page":"D1448","DOI":"10.1093\/nar\/gkab949","article-title":"PCMDB: a curated and comprehensive resource of plant cell markers","volume":"50","author":"Jin","year":"2022","journal-title":"Nucleic Acids Research"},{"key":"2026091519522645500_erag287-B22","doi-asserted-by":"crossref","first-page":"1065","DOI":"10.2307\/3869885","article-title":"Roles of the GLABROUS1 and TRANSPARENT TESTA GLABRA genes in Arabidopsis trichome development","volume":"6","author":"Larkin","year":"1994","journal-title":"The Plant Cell"},{"key":"2026091519522645500_erag287-B23","doi-asserted-by":"crossref","first-page":"694","DOI":"10.1016\/j.devcel.2023.03.004","article-title":"Single-nucleus sequencing deciphers developmental trajectories in rice pistils","volume":"58","author":"Li","year":"2023","journal-title":"Developmental Cell"},{"key":"2026091519522645500_erag287-B24","doi-asserted-by":"crossref","first-page":"2515","DOI":"10.1093\/jxb\/erv046","article-title":"The identification of Cucumis sativus Glabrous 1 (CsGL1) required for the formation of trichomes uncovers a novel function for the homeodomain-leucine zipper I gene","volume":"66","author":"Li","year":"2015","journal-title":"Journal of Experimental Botany"},{"key":"2026091519522645500_erag287-B25","article-title":"Single-cell transcriptome atlas and regulatory dynamics in developing cotton anthers","volume":"11","author":"Li","year":"2024","journal-title":"Advanced Science"},{"key":"2026091519522645500_erag287-B26","doi-asserted-by":"crossref","first-page":"1310","DOI":"10.1016\/j.molp.2022.05.013","article-title":"Allele-aware chromosome-level genome assembly of Artemisia annua reveals the correlation between ADS expansion and artemisinin yield","volume":"15","author":"Liao","year":"2022","journal-title":"Molecular Plant"},{"key":"2026091519522645500_erag287-B27","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1007\/s11240-016-1015-4","article-title":"Characterization of a trichome-specific promoter of the aldehyde dehydrogenase 1 (ALDH1) gene in Artemisia annua","volume":"126","author":"Liu","year":"2016","journal-title":"Plant Cell, Tissue and Organ Culture"},{"key":"2026091519522645500_erag287-B28","doi-asserted-by":"crossref","first-page":"1734","DOI":"10.1038\/s41477-023-01524-8","article-title":"Single-nucleus transcriptomes reveal spatiotemporal symbiotic perception and early response in Medicago","volume":"9","author":"Liu","year":"2023","journal-title":"Nature Plants"},{"key":"2026091519522645500_erag287-B29","doi-asserted-by":"crossref","first-page":"428","DOI":"10.1038\/s41588-020-0604-7","article-title":"Stepwise selection on homeologous PRR genes controlling flowering and maturity during soybean domestication","volume":"52","author":"Lu","year":"2020","journal-title":"Nature Genetics"},{"key":"2026091519522645500_erag287-B30","doi-asserted-by":"crossref","first-page":"1191","DOI":"10.1111\/nph.12207","article-title":"AaORA, a trichome-specific AP2\/ERF transcription factor of Artemisia annua, is a positive regulator in the artemisinin biosynthetic pathway and in disease resistance to Botrytis cinerea","volume":"198","author":"Lu","year":"2013","journal-title":"New Phytologist"},{"key":"2026091519522645500_erag287-B31","doi-asserted-by":"crossref","first-page":"1212","DOI":"10.1111\/jipb.13258","article-title":"The transcription factors TLR1 and TLR2 negatively regulate trichome density and artemisinin levels in Artemisia annua","volume":"64","author":"Lv","year":"2022","journal-title":"Journal of Integrative Plant Biology"},{"key":"2026091519522645500_erag287-B32","doi-asserted-by":"crossref","first-page":"307","DOI":"10.4103\/wjtcm.wjtcm_20_21","article-title":"Phytohormones jasmonic acid, salicylic acid, gibberellins, and abscisic acid are key mediators of plant secondary metabolites","volume":"7","author":"Lv","year":"2021","journal-title":"World Journal of Traditional Chinese Medicine"},{"key":"2026091519522645500_erag287-B33","doi-asserted-by":"crossref","first-page":"2146","DOI":"10.1093\/pcp\/pcp149","article-title":"Isolation and characterization of AaWRKY1, an Artemisia annua transcription factor that regulates the amorpha-4,11-diene synthase gene, a key gene of artemisinin biosynthesis","volume":"50","author":"Ma","year":"2009","journal-title":"Plant & Cell Physiology"},{"key":"2026091519522645500_erag287-B34","doi-asserted-by":"crossref","first-page":"1624","DOI":"10.1104\/pp.15.01309","article-title":"TEMPRANILLO reveals the mesophyll as crucial for epidermal trichome formation","volume":"170","author":"Mat\u00edas-Hern\u00e1ndez","year":"2016","journal-title":"Plant Physiology"},{"key":"2026091519522645500_erag287-B35","doi-asserted-by":"crossref","first-page":"473","DOI":"10.1111\/j.1399-3054.1962.tb08052.x","article-title":"A revised medium for rapid growth and bio assays with tobacco tissue cultures","volume":"15","author":"Murashige","year":"1962","journal-title":"Physiologia Plantarum"},{"key":"2026091519522645500_erag287-B36","doi-asserted-by":"crossref","first-page":"1123","DOI":"10.1016\/j.phytochem.2009.07.009","article-title":"Localization of enzymes of artemisinin biosynthesis to the apical cells of glandular secretory trichomes of Artemisia annua L","volume":"70","author":"Olsson","year":"2009","journal-title":"Phytochemistry"},{"key":"2026091519522645500_erag287-B37","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1016\/0092-8674(91)90523-2","article-title":"A myb gene required for leaf trichome differentiation in Arabidopsis is expressed in stipules","volume":"67","author":"Oppenheimer","year":"1991","journal-title":"Cell"},{"key":"2026091519522645500_erag287-B38","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1186\/s12936-021-03987-6","article-title":"Prevalence of potential mediators of artemisinin resistance in African isolates of Plasmodium falciparum","volume":"20","author":"Owoloye","year":"2021","journal-title":"Malaria Journal"},{"key":"2026091519522645500_erag287-B39","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1186\/s12870-015-0693-0","article-title":"The loss-of-function GLABROUS 3 mutation in cucumber is due to LTR-retrotransposon insertion in a class IV HD-ZIP transcription factor gene CsGL3 that is epistatic over CsGL1","volume":"15","author":"Pan","year":"2015","journal-title":"BMC Plant Biology"},{"key":"2026091519522645500_erag287-B40","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":"2026091519522645500_erag287-B41","doi-asserted-by":"crossref","first-page":"1795","DOI":"10.1105\/tpc.111.083261","article-title":"The jasmonate-ZIM-domain proteins interact with the WD-repeat\/bHLH\/MYB complexes to regulate jasmonate-mediated anthocyanin accumulation and trichome initiation in Arabidopsis thaliana","volume":"23","author":"Qi","year":"2011","journal-title":"The Plant Cell"},{"key":"2026091519522645500_erag287-B42","doi-asserted-by":"crossref","first-page":"2372","DOI":"10.1111\/pbi.13918","article-title":"Single-cell RNA-seq reveals fate determination control of an individual fibre cell initiation in cotton (Gossypium hirsutum)","volume":"20","author":"Qin","year":"2022","journal-title":"Plant Biotechnology Journal"},{"key":"2026091519522645500_erag287-B43","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1387\/ijdb.051983ss","article-title":"Epidermal differentiation: trichomes in Arabidopsis as a model system","volume":"49","author":"Schellmann","year":"2005","journal-title":"The International Journal of Developmental Biology"},{"key":"2026091519522645500_erag287-B44","doi-asserted-by":"crossref","first-page":"5036","DOI":"10.1093\/emboj\/cdf524","article-title":"TRIPTYCHON and CAPRICE mediate lateral inhibition during trichome and root hair patterning in Arabidopsis","volume":"21","author":"Schellmann","year":"2002","journal-title":"The EMBO Journal"},{"key":"2026091519522645500_erag287-B45","doi-asserted-by":"crossref","first-page":"5519","DOI":"10.1038\/ncomms6519","article-title":"Control of cotton fibre elongation by a homeodomain transcription factor GhHOX3","volume":"5","author":"Shan","year":"2014","journal-title":"Nature Communications"},{"key":"2026091519522645500_erag287-B46","doi-asserted-by":"crossref","first-page":"1269","DOI":"10.1111\/nph.13874","article-title":"The jasmonate-responsive AaMYC2 transcription factor positively regulates artemisinin biosynthesis in Artemisia annua","volume":"210","author":"Shen","year":"2016","journal-title":"New Phytologist"},{"key":"2026091519522645500_erag287-B47","doi-asserted-by":"crossref","first-page":"776","DOI":"10.1016\/j.molp.2018.03.015","article-title":"The genome of Artemisia annua provides insight into the evolution of Asteraceae family and artemisinin biosynthesis","volume":"11","author":"Shen","year":"2018","journal-title":"Molecular Plant"},{"key":"2026091519522645500_erag287-B48","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1111\/nph.14789","article-title":"The roles of AaMIXTA1 in regulating the initiation of glandular trichomes and cuticle biosynthesis in Artemisia annua","volume":"217","author":"Shi","year":"2018","journal-title":"New Phytologist"},{"key":"2026091519522645500_erag287-B49","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1186\/1471-2229-13-220","article-title":"Differential transcriptome analysis of glandular and filamentous trichomes in Artemisia annua","volume":"13","author":"Soetaert","year":"2013","journal-title":"BMC Plant Biology"},{"key":"2026091519522645500_erag287-B50","doi-asserted-by":"crossref","first-page":"694","DOI":"10.1016\/j.molp.2023.02.005","article-title":"Single-cell transcriptomic analysis reveals the developmental trajectory and transcriptional regulatory networks of pigment glands in Gossypium bickii","volume":"16","author":"Sun","year":"2023","journal-title":"Molecular Plant"},{"key":"2026091519522645500_erag287-B51","doi-asserted-by":"crossref","first-page":"1396","DOI":"10.1016\/j.molp.2015.04.002","article-title":"TRICHOME AND ARTEMISININ REGULATOR 1 is required for trichome development and artemisinin biosynthesis in Artemisia annua","volume":"8","author":"Tan","year":"2015","journal-title":"Molecular Plant"},{"key":"2026091519522645500_erag287-B52","doi-asserted-by":"crossref","first-page":"898","DOI":"10.1093\/plphys\/kiab489","article-title":"Single-cell transcriptomics sheds light on the identity and metabolism of developing leaf cells","volume":"188","author":"Tenorio Berr\u00edo","year":"2022","journal-title":"Plant Physiology"},{"key":"2026091519522645500_erag287-B53","doi-asserted-by":"crossref","first-page":"3499","DOI":"10.1093\/jxb\/eraa104","article-title":"The transcription factor MML4_D12 regulates fiber development through interplay with the WD40-repeat protein WDR in cotton","volume":"71","author":"Tian","year":"2020","journal-title":"Journal of Experimental Botany"},{"key":"2026091519522645500_erag287-B54","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1111\/plb.70131","article-title":"Single-cell RNA sequencing methodology, analysis, applications, and future directions with special focus on cotton","volume":"28","author":"Verma","year":"2026","journal-title":"Plant Biology"},{"key":"2026091519522645500_erag287-B55","doi-asserted-by":"crossref","first-page":"785","DOI":"10.1111\/j.1365-313X.2010.04464.x","article-title":"GhMYB25-like: a key factor in early cotton fibre development","volume":"65","author":"Walford","year":"2011","journal-title":"The Plant Journal"},{"key":"2026091519522645500_erag287-B56","doi-asserted-by":"crossref","first-page":"1337","DOI":"10.1105\/tpc.11.7.1337","article-title":"The TRANSPARENT TESTA GLABRA1 locus, which regulates trichome differentiation and anthocyanin biosynthesis in Arabidopsis, encodes a WD40 repeat protein","volume":"11","author":"Walker","year":"1999","journal-title":"The Plant Cell"},{"key":"2026091519522645500_erag287-B57","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1016\/j.ymben.2016.07.004","article-title":"Transient production of artemisinin in Nicotiana benthamiana is boosted by a specific lipid transfer protein from A. annua","volume":"38","author":"Wang","year":"2016","journal-title":"Metabolic Engineering"},{"key":"2026091519522645500_erag287-B58","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1186\/s13059-023-02886-0","article-title":"Cell-specific clock-controlled gene expression program regulates rhythmic fiber cell growth in cotton","volume":"24","author":"Wang","year":"2023","journal-title":"Genome Biology"},{"key":"2026091519522645500_erag287-B59","doi-asserted-by":"crossref","first-page":"619","DOI":"10.4236\/ajps.2011.24073","article-title":"Roles of the GLABROUS1 and TRANSPARENT TESTA GLABRA genes in Arabidopsis trichome development","volume":"2","author":"Wang","year":"2011","journal-title":"Americal Journal of Plant Sciences"},{"key":"2026091519522645500_erag287-B60","doi-asserted-by":"crossref","first-page":"2323","DOI":"10.1105\/tpc.104.024844","article-title":"Control of plant trichome development by a cotton fiber MYB gene","volume":"16","author":"Wang","year":"2004","journal-title":"The Plant Cell"},{"key":"2026091519522645500_erag287-B61","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1186\/1471-2164-10-465","article-title":"Global characterization of Artemisia annua glandular trichome transcriptome using 454 pyrosequencing","volume":"10","author":"Wang","year":"2009","journal-title":"BMC Genomics"},{"key":"2026091519522645500_erag287-B62","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1007\/s00425-021-03676-3","article-title":"Enhancing artemisinin content in and delivery from Artemisia annua: a review of alternative, classical, and transgenic approaches","volume":"254","author":"Wani","year":"2021","journal-title":"Planta"},{"key":"2026091519522645500_erag287-B63","doi-asserted-by":"crossref","DOI":"10.1016\/j.jhazmat.2023.130899","article-title":"Insights into strigolactone (GR24) mediated regulation of cadmium-induced changes and ROS metabolism in Artemisia annua","volume":"448","author":"Wani","year":"2023","journal-title":"Journal of Hazardous Materials"},{"key":"2026091519522645500_erag287-B64","doi-asserted-by":"crossref","DOI":"10.1016\/j.plantsci.2023.111754","article-title":"Nitric oxide induces antioxidant machinery, PSII functioning and artemisinin biosynthesis in Artemisia annua under cadmium stress","volume":"334","author":"Wani","year":"2023","journal-title":"Plant Science"},{"key":"2026091519522645500_erag287-B65","doi-asserted-by":"crossref","first-page":"278","DOI":"10.1016\/j.devcel.2023.01.009","article-title":"A HD-ZIP transcription factor specifies fates of multicellular trichomes via dosage-dependent mechanisms in tomato","volume":"58","author":"Wu","year":"2023","journal-title":"Developmental Cell"},{"key":"2026091519522645500_erag287-B66","doi-asserted-by":"crossref","first-page":"2375","DOI":"10.1093\/plcell\/koae077","article-title":"A gradient of the HD-Zip regulator woolly regulates multicellular trichome morphogenesis in tomato","volume":"36","author":"Wu","year":"2024","journal-title":"The Plant Cell"},{"key":"2026091519522645500_erag287-B67","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1104\/pp.110.161646","article-title":"The glabra1 mutation affects cuticle formation and plant responses to microbes","volume":"154","author":"Xia","year":"2010","journal-title":"Plant Physiology"},{"key":"2026091519522645500_erag287-B68","doi-asserted-by":"crossref","first-page":"2050","DOI":"10.1111\/nph.17514","article-title":"An HD-ZIP-MYB complex regulates glandular secretory trichome initiation in Artemisia annua","volume":"231","author":"Xie","year":"2021","journal-title":"New Phytologist"},{"key":"2026091519522645500_erag287-B69","doi-asserted-by":"crossref","first-page":"1691","DOI":"10.1093\/jxb\/eraa523","article-title":"The WRKY transcription factor AaGSW2 promotes glandular trichome initiation in Artemisia annua","volume":"72","author":"Xie","year":"2021","journal-title":"Journal of Experimental Botany"},{"key":"2026091519522645500_erag287-B70","doi-asserted-by":"crossref","first-page":"1145","DOI":"10.1111\/nph.14205","article-title":"HOMEODOMAIN PROTEIN 1 is required for jasmonate-mediated glandular trichome initiation in Artemisia annua","volume":"213","author":"Yan","year":"2017","journal-title":"New Phytologist"},{"key":"2026091519522645500_erag287-B71","doi-asserted-by":"crossref","first-page":"567","DOI":"10.1111\/nph.15005","article-title":"A novel HD-ZIP IV\/MIXTA complex promotes glandular trichome initiation and cuticle development in Artemisia annua","volume":"218","author":"Yan","year":"2018","journal-title":"New Phytologist"},{"key":"2026091519522645500_erag287-B72","doi-asserted-by":"crossref","first-page":"11836","DOI":"10.1073\/pnas.1100532108","article-title":"A regulatory gene induces trichome formation and embryo lethality in tomato","volume":"108","author":"Yang","year":"2011","journal-title":"Proceedings of the National Academy of Sciences, USA"},{"key":"2026091519522645500_erag287-B73","doi-asserted-by":"crossref","first-page":"3320","DOI":"10.1105\/tpc.112.101014","article-title":"Gibberellin regulates the Arabidopsis floral transition through miR156-targeted SQUAMOSA promoter binding-like transcription factors","volume":"24","author":"Yu","year":"2012","journal-title":"The Plant Cell"},{"key":"2026091519522645500_erag287-B74","doi-asserted-by":"crossref","first-page":"885","DOI":"10.1111\/nph.18567","article-title":"AabHLH113 integrates jasmonic acid and abscisic acid signaling to positively regulate artemisinin biosynthesis in Artemisia annua","volume":"237","author":"Yuan","year":"2023","journal-title":"New Phytologist"},{"key":"2026091519522645500_erag287-B75","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1111\/pbi.13448","article-title":"R2r3 MYB-dependent auxin signalling regulates trichome formation, and increased trichome density confers spider mite tolerance on tomato","volume":"19","author":"Yuan","year":"2021","journal-title":"Plant Biotechnology Journal"},{"key":"2026091519522645500_erag287-B76","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1038\/s41467-023-44369-7","article-title":"Decoding the gene regulatory network of endosperm differentiation in maize","volume":"15","author":"Yuan","year":"2024","journal-title":"Nature Communications"},{"key":"2026091519522645500_erag287-B77","doi-asserted-by":"crossref","DOI":"10.1016\/j.indcrop.2019.111476","article-title":"Overexpression of AaPIF3 promotes artemisinin production in Artemisia annua","volume":"138","author":"Zhang","year":"2019","journal-title":"Industrial Crops and Products"},{"key":"2026091519522645500_erag287-B78","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1105\/tpc.114.135186","article-title":"An intrinsic microRNA timer regulates progressive decline in shoot regenerative capacity in plants","volume":"27","author":"Zhang","year":"2015","journal-title":"The Plant Cell"}],"container-title":["Journal of Experimental Botany"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/jxb\/advance-article-pdf\/doi\/10.1093\/jxb\/erag287\/68540071\/erag287.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jxb\/article-pdf\/77\/17\/5715\/68540071\/erag287.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jxb\/article-pdf\/77\/17\/5715\/68540071\/erag287.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T23:52:46Z","timestamp":1789516366000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/jxb\/article\/77\/17\/5715\/8709591"}},"subtitle":[],"editor":[{"given":"John","family":"Lunn","sequence":"additional","affiliation":[{"name":"Max-Planck-Institut f\u00fcr Molekulare Pflanzenphysiologie, Germany"}],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,6,17]]},"references-count":78,"journal-issue":{"issue":"17","published-online":{"date-parts":[[2026,6,17]]},"published-print":{"date-parts":[[2026,9,16]]}},"URL":"https:\/\/doi.org\/10.1093\/jxb\/erag287","relation":{},"ISSN":["0022-0957","1460-2431"],"issn-type":[{"value":"0022-0957","type":"print"},{"value":"1460-2431","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,9,16]]},"published":{"date-parts":[[2026,6,17]]}}}