Clinical MedicineHematologyImmunology Open Access | 10.1172/JCI173096

Gianluca Storci,1 Francesco De Felice,1,2 Francesca Ricci,1 Spartaco Santi,3,4 Daria Messelodi,1 Salvatore Nicola Bertuccio,1 Noemi Laprovitera,1 Michele Dicataldo,1,2 Lucrezia Rossini,2 Serena De Matteis,1 Beatrice Casadei,1 Francesca Vaglio,1 Margherita Ursi,1,2 Francesco Barbato,1,2 Marcello Roberto,1,2 Maria Guarino,1 Gian Maria Asioli,1 Mario Arpinati,1 Pietro Cortelli,5,6 Enrico Maffini,1 Enrica Tomassini,1 Marta Tassoni,2 Carola Cavallo,7 Francesco Iannotta,1 Maria Naddeo,1,2 Pier Luigi Tazzari,1 Elisa Dan,1 Cinzia Pellegrini,1 Serafina Guadagnuolo,1 Matteo Carella,1,2 Barbara Sinigaglia,1 Chiara Pirazzini,2 Caterina Severi,8 Paolo Garagnani,1,2 Katarzyna Malgorzata Kwiatkowska,2 Manuela Ferracin,1,2 Pier Luigi Zinzani,2,9 Massimiliano Bonafè,1,2 and Francesca Bonifazi1

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

Find articles by De Felice, F. in: JCI | PubMed | Google Scholar

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

Find articles by Ricci, F. in: JCI | PubMed | Google Scholar

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

Find articles by Santi, S. in: JCI | PubMed | Google Scholar |

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

Find articles by Messelodi, D. in: JCI | PubMed | Google Scholar

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

Find articles by Laprovitera, N. in: JCI | PubMed | Google Scholar

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

Find articles by Dicataldo, M. in: JCI | PubMed | Google Scholar

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

Find articles by Rossini, L. in: JCI | PubMed | Google Scholar

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

Find articles by De Matteis, S. in: JCI | PubMed | Google Scholar

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

Find articles by Casadei, B. in: JCI | PubMed | Google Scholar

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

Find articles by Roberto, M. in: JCI | PubMed | Google Scholar

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

Find articles by Maffini, E. in: JCI | PubMed | Google Scholar

1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria di Bologna, Università di Bologna, Via San Giacomo 14, 40134 Bologna, Italy. Phone: 39.51.2094748; Email: massimiliano.bonafe@unibo.it.

Authorship note: GS and FDF contributed equally to this work and are co–first authors. MB and F Bonifazi contributed equally to this work and are co–senior authors.

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1IRCCS Azienda Ospedaliero-Universitaria di Bologna, Bologna, Italy.

2Dipartimento di Scienze Mediche e Chirurgiche, Università di Bologna, Bologna, Italy.

3IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

4Institute of Molecular Genetics, National Research Council of Italy, Bologna, Italy.

5Department of Biomedical and Neuromotor Sciences, Bellaria Hospital, Università di Bologna, Bologna, Italy.

6IRCCS Institute of Neurological Sciences of Bologna, Bellaria Hospital, Bologna, Italy.

7Laboratory Ramses, Research & Innovation Technology Department, IRCCS Istituto Ortopedico Rizzoli, Bologna, Italy.

8Abbelight, Cachan, France.

9IRCCS Azienda Ospedaliero–Università di Bologna, Istituto di Ematologia “Seràgnoli,” Bologna, Italy.

Address correspondence to: Massimiliano Bonafè, IRCCS Azienda Ospedaliero–Universitaria