Genah, Shirley
Pellegrino, Marsha
Giansanti, Manuela
Mancusi, Angelica https://orcid.org/0009-0008-0150-9825
Taviani, Alessio
Cardinale, Antonella
Rosichini, Marco
Flamini, Sara
Catanoso, Maria Luigia
Giorda, Ezio https://orcid.org/0000-0002-5875-832X
Volpe, Gabriele
Piccione, Michela
Nazio, Francesca
Dudakov, Jarrod A. https://orcid.org/0000-0001-6220-5632
van den Brink, Marcel R. M. https://orcid.org/0000-0003-0696-4401
de Billy, Emmanuel
Locatelli, Franco https://orcid.org/0000-0002-7976-3654
Velardi, Enrico https://orcid.org/0000-0001-8383-0453
Funding for this research was provided by:
Ministero della Salute (GR-2019-12369231)
Ministero della Salute (RF-2016-02364388)
Be The Match Foundation
Associazione Italiana per la Ricerca sul Cancro (29932)
Associazione Italiana per la Ricerca sul Cancro (27019)
Associazione Italiana per la Ricerca sul Cancro (21147)
Rally Foundation
Amy Stelzer Manasevit Research Program
Ministero dell'Istruzione, dell'Università e della Ricerca (PRIN ID 2017 WC8499_004)
Article History
Received: 6 June 2025
Revised: 11 February 2026
Accepted: 10 March 2026
First Online: 26 March 2026
Competing interests
: GS is supported by a FIRC-AIRC fellowship for Italy. RM was supported by an AIRC Fellowship for Abroad. VE was supported by grants from the Rally Foundation Investigator Award, Amy Stelzer Manasevit Research Program, the Italian Association for Cancer Research, and by the Italian Ministry of Health with “Current Research funds. LF was supported by grants from AIRC (Special Program Metastatic disease: the key unmet need in oncology 5 per mille 2018 Project Code 21147 and Accelerator Award 2017 INCAR); Ministero dell’Istruzione, dell’Università e della Ricerca, PRIN ID 2017 WC8499_004; Ministero della Salute, RF-2016-02364388. EdB received funding from Research (PRIN PNRR P2022JLHZZ), Ministero della Salute (GR-2019-12369231), by MIA Neri Foundation and by Heal Foundation. FN lab is supported by the “Associazione Italiana Ricerca sul Cancro” (My First Airc Grant -Ref. 27019), by “CureSearch Young Investigator Awards in Pediatric Oncology Drug Development”, by “Fondazione Roche per la Ricerca Indipendente”, by the Italian Ministry of University and Research (PRIN PNRR P2022JLHZZ), Ministero della Salute (GR-2019-12369231) and by MIA Neri Foundation. EV has received research funding from ToleranceBio for projects outside the submitted work. FL reports personal fees from Amgen, personal fees from Novartis, other from Bellicum Pharmaceutical, other from Neovii, personal fees from Miltenyi, personal fees from Medac, personal fees from Jazz Pharmaceutical, personal fees from Takeda, outside the submitted work. A patent application entitled “FOXN1-activating compounds and their use in thymic regeneration” has been filed (PCT/IT2025/050304).
: All animals were purchased from Charles River and maintained in the Plaisant Castel Romano animal facility in Rome. Mice experiments were conducted in compliance with the EU and national ethical requirements and were approved by the Italian Health Ministry. Human thymic samples for hTEC culture were obtained from pediatric patients undergoing corrective cardiac surgery. All samples were anonymously collected in accordance with local ethical guidelines, written informed consent was obtained, and the protocol was approved by the Ethics Committee of OPBG Hospital.