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Senescent macrophages are functionally heterogeneous: canonical p16-high macrophages promote tumorigenesis or, in other contexts, disease tolerance, whereas we previously identified a distinct p21-high, p16-low senotype that drives metabolic dysfunction-associated steatotic liver disease (MASLD). The molecular basis of this senotype has remained undefined. Using genetic and multi-omic approaches, we show that a p53-p21-dependent program actively represses p16 and is required for senescent macrophage viability. We identify Cyclin D2 as a non-canonical downstream effector that redistributes from the nucleus to mitochondria and lipid droplets, where it partners with MIC60 to drive metabolic reprogramming and modulates AKT1-mTORC1 signaling that sustains the SASP. Cyclin D2-p21 double-positive macrophages accumulate with aging and MASLD in mice and in human liver cirrhosis, and can be selectively depleted by senolytic treatment. Together, these findings define a druggable p53-p21-Cyclin D2 axis that specifies macrophage senotype.<\/jats:p>\n                <jats:sec>\n                  <jats:title>Highlights<\/jats:title>\n                  <jats:list list-type=\"simple\">\n                    <jats:list-item>\n                      <jats:p>\u25cf A p53\u2013p21-dependent program drives macrophage senescence while actively repressing p16<\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\u25cf p16 repression promotes cell survival during acute genotoxic stress, but is dispensable for cell-cycle arrest and core senescence features in macrophages<\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\u25cf Cyclin D2 is a non-canonical p53\u2013p21 effector that relocalizes from nucleus to the cytosol and mitochondria<\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\u25cf Cyclin D2 engages the AKT1-mTORC1 pathway and interacts with MIC60 to drive metabolic reprogramming<\/jats:p>\n                    <\/jats:list-item>\n                    <jats:list-item>\n                      <jats:p>\u25cf Cyclin D2+p21+ macrophages are abundant in aging, MASLD, and human liver cirrhosis, and can be targeted by senolytics<\/jats:p>\n                    <\/jats:list-item>\n                  <\/jats:list>\n                <\/jats:sec>","DOI":"10.64898\/2026.09.22.753597","type":"posted-content","created":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T16:45:21Z","timestamp":1790613921000},"source":"Crossref","is-referenced-by-count":0,"title":["The p53\u2013p21\u2013Cyclin D2 regulatory axis drives metabolic reprogramming and a distinct senescent macrophage senotype during aging and 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