This project has received funding from the Giovanni Armenise-Harvard Foundation, USA (Career Development Award 2016, to L.T.), Pezcoller Foundation, CARITRO (to L.T.), My First AIRC Grant, Italy (Project Code: 19921 to L.T.) and EMBO YIP small grant to L.T. F.A has received support by the European Union’s Horizon 2020 research and innovation programme under the Marie Skłodowska Curie grant agreement no. 844677 (to F.A.) and by the University of Trento under the action MSCA+1. F.A. was also supported by Fondazione Umberto Veronesi postdoctoral fellowship (il Dono di Rossana). M.G. has been supported by a doctoral fellowship funded by Fondazione Pezcoller. We thank S. Robbiati (MOF facility), M. Pancher (HTS facility), M. Cardano (Cell Technology facility), G. Scarduelli and M. Roccuzzo (AICF) of the University of Trento.