Cellular insults, such as oxidative stress or DNA damage, lead to nucleolar stress, namely disruption of nucleolar function and structure, stabilization of p53 and p53-dependent cell cycle arrest, senescence and apoptosis, reinforcing the central role of p53 in surveying cellular health.9, 10 This tight link between RNA-polymerase I (Pol I) activity, nucleolar integrity and p53-mediated damage con