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Characterizing mutants with impaired riboflavin metabolism can help clarify the role of riboflavin in plant development. Here, we characterized a rice (\n                    <jats:italic>Oryza sativa<\/jats:italic>\n                    )\n                    <jats:italic>white and lesion\u2010mimic<\/jats:italic>\n                    (\n                    <jats:italic>wll1<\/jats:italic>\n                    ) mutant, which displays a lesion\u2010mimic phenotype with white leaves, chlorophyll loss, chloroplast defects, excess reactive oxygen species (ROS) accumulation, decreased photosystem protein levels, changes in expression of chloroplast development and photosynthesis genes, and cell death. Map\u2010based cloning and complementation test revealed that\n                    <jats:italic>WLL1<\/jats:italic>\n                    encodes lumazine synthase, which participates in riboflavin biosynthesis. Indeed, the\n                    <jats:italic>wll1<\/jats:italic>\n                    mutant showed riboflavin deficiency, and application of FAD rescued the\n                    <jats:italic>wll1<\/jats:italic>\n                    phenotype. In addition, transcriptome analysis showed that cytokinin metabolism was significantly affected in\n                    <jats:italic>wll1<\/jats:italic>\n                    mutant, which had increased cytokinin and \u03b4\u2010aminolevulinic acid contents. Furthermore, WLL1 and riboflavin synthase (RS) formed a complex, and the\n                    <jats:italic>rs<\/jats:italic>\n                    mutant had a similar phenotype to the\n                    <jats:italic>wll1<\/jats:italic>\n                    mutant. Taken together, our findings revealed that\n                    <jats:italic>WLL1<\/jats:italic>\n                    and\n                    <jats:italic>RS<\/jats:italic>\n                    play pivotal roles in riboflavin biosynthesis, which is necessary for ROS balance and chloroplast development in rice.\n                  <\/jats:p>","DOI":"10.1111\/tpj.15537","type":"journal-article","created":{"date-parts":[[2021,10,10]],"date-time":"2021-10-10T13:26:35Z","timestamp":1633872395000},"page":"1690-1703","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":24,"title":["<i>WHITE AND LESION\u2010MIMIC LEAF1<\/i>\n                    , encoding a lumazine synthase, affects reactive oxygen species balance and chloroplast development in rice"],"prefix":"10.1111","volume":"108","author":[{"given":"Haitao","family":"Hu","sequence":"first","affiliation":[{"name":"State Key Laboratory of Rice Biology China National Rice Research Institute Hangzhou 310006 China"},{"name":"College of Chemistry and Life Sciences Zhejiang Normal University Jinhua 321004 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1140-3633","authenticated-orcid":false,"given":"Deyong","family":"Ren","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Rice Biology China National Rice Research Institute Hangzhou 310006 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiang","family":"Hu","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Rice Biology China National Rice Research Institute Hangzhou 310006 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongzhen","family":"Jiang","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Rice Biology China National Rice Research Institute Hangzhou 310006 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ping","family":"Chen","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Rice Biology China National Rice Research Institute Hangzhou 310006 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dali","family":"Zeng","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Rice Biology China National Rice Research Institute Hangzhou 310006 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qian","family":"Qian","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Rice Biology China National Rice Research Institute Hangzhou 310006 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Longbiao","family":"Guo","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Rice Biology China National Rice Research Institute Hangzhou 310006 China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2021,10,27]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.17467"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.1039\/C8CS00154E"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.1002\/jimd.12058"},{"key":"e_1_2_9_5_1","first-page":"365","article-title":"Porphyrin and heme metabolism and the porphyrias","volume":"3","author":"Bonkovsky H.L.","year":"2012","journal-title":"Comprehensive Physiology"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbabio.2015.05.007"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10725-018-0403-7"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.13299"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.16.00640"},{"key":"e_1_2_9_10_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erv132"},{"key":"e_1_2_9_11_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.15448"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.15079"},{"key":"e_1_2_9_13_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erz268"},{"key":"e_1_2_9_14_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.phytochem.2021.112822"},{"key":"e_1_2_9_15_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00425-006-0249-5"},{"key":"e_1_2_9_16_1","doi-asserted-by":"publisher","DOI":"10.1016\/B978-0-12-386479-6.00003-2"},{"key":"e_1_2_9_17_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.17127"},{"key":"e_1_2_9_18_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.105.033589"},{"key":"e_1_2_9_19_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-313X.2004.02262.x"},{"key":"e_1_2_9_20_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.20.00445"},{"key":"e_1_2_9_21_1","doi-asserted-by":"publisher","DOI":"10.1007\/s11103-011-9846-1"},{"key":"e_1_2_9_22_1","unstructured":"Hiltunen H.(2016)Functional Analysis of RIBA the Introductory Enzyme for Riboflavin Biosynthesis. 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