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Helsinki Institute of Life Science, Biomedicum Helsinki, University of Helsinki, Helsinki, Finland; Wihuri Research Institute, Biomedicum Helsinki, University of Helsinki, Helsinki, Finland; Stem Cells and Metabolism Research Program, Faculty of Medicine, University of Helsinki, Helsinki, Finland; Max Planck Institute for Biology of Ageing, Cologne, Germany; Cologne Excellence Cluster for Stress Responses in Ageing-associated diseases (CECAD), University of Cologne, Cologne, Germany. Electronic address: