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The\n                          <jats:italic>Polima<\/jats:italic>\n                          (\n                          <jats:italic>pol<\/jats:italic>\n                          ) CMS system is associated with the mitochondrial chimeric gene\n                          <jats:italic>orf224<\/jats:italic>\n                          , yet how it disrupts early anther differentiation and leads to sterility remains unclear.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          We combined cytological analyses, ultrastructural observations, transcriptomics, proteomics, and metabolomics to examine young anther development in\n                          <jats:italic>pol<\/jats:italic>\n                          CMS, maintainer, and restorer lines, with particular focus on mitochondrial function, metabolism, and hormone homeostasis.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          In\n                          <jats:italic>pol<\/jats:italic>\n                          CMS, L2\u2010derived anther cells failed to differentiate into the middle layer, tapetum, and microspore mother cells, with development arrested at stage S5. The ORF224 protein preferentially accumulated in young anthers and exhibited cytotoxicity, causing mitochondrial dysfunction, reduced ATP production, and activation of mitochondrial retrograde signaling. These changes induced extensive metabolic reprogramming, including ectopic activation of photosynthesis\u2010like reaction, starch accumulation, elevated ROS, and a pronounced increase in abscisic acid (ABA).\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          Our results demonstrate that\n                          <jats:italic>pol<\/jats:italic>\n                          CMS sterility results from ORF224\u2010mediated mitochondrial stress that reprograms anther cell fate from reproductive to somatic development, with ABA reinforcing developmental arrest, while programmed cell death occurs as a late downstream consequence.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                    <\/jats:list>\n                  <\/jats:p>","DOI":"10.1111\/nph.71462","type":"journal-article","created":{"date-parts":[[2026,7,27]],"date-time":"2026-07-27T08:23:35Z","timestamp":1785140615000},"page":"3384-3400","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Mitochondrial gene\n                    <i>orf224<\/i>\n                    triggers retrograde signaling and reprogramming of anther cell fate in\n                    <i>Brassica napus Polima<\/i>\n                    <scp>CMS<\/scp>"],"prefix":"10.1111","volume":"251","author":[{"ORCID":"https:\/\/orcid.org\/0009-0001-8195-5591","authenticated-orcid":false,"given":"Yilei","family":"Zhang","sequence":"first","affiliation":[{"name":"State Key Laboratory of Biocatalysis and Enzyme Engineering, School of Life Sciences Hubei University  Wuhan 430070 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