Reactivation of telomerase in cultured human cells extends their replicative life span beyond the Hayflick limit. How telomere shortening triggers cell senescence and whether it contributes to aging in vivo are under investigation. Studies in yeast have revealed another site critical to cellular aging: the nucleolus. The accumulation of ribosomal DNA circles is a cause of aging in this organism. The possible relevance of this mechanism to human aging is also being considered.
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Johnson, F. B., Marciniak, R. A., & Guarente, L. (1998). Telomeres, the nucleolus and aging. Current Opinion in Cell Biology, 10(3), 332–338. https://doi.org/10.1016/S0955-0674(98)80008-2