Role of glutamine synthetase in angiogenesis 1 beyond glutamine synthesis 2 3

3 Guy Eelen*, Charlotte Dubois*, Anna Rita Cantelmo, Jermaine Goveia, Ulrike Brün4 ing, Michael DeRan, Gopala Jarugumilli, Jos van Rijssel, Giorgio Saladino, Federico 5 Comitani, Annalisa Zecchin, Susana Rocha, Hongling Huang, Saar Vandekeere, Jo6 anna Kalucka, Christian Lange, Francisco Morales-Rodriguez, Bert Cruys, Lucas 7 Treps, Leanne Ramer, Stefan Vinckier, Katleen Brepoels, Sabine Wyns, Joris 8 Souffreau, Luc Schoonjans, Wouter H. Lamers, Yi Wu, Jurgen Haustraete, Johan 9 Hofkens, Sandra Liekens, Richard Cubbon, Bart Ghesquière, Mieke Dewerchin, 10 Francesco L. Gervasio, Jaap D. van Buul, Xu Wu & Peter Carmeliet 11 12 (1) Center for Cancer Biology, University of Leuven, Leuven, B-3000, Belgium; (2) Center for Cancer Biolo13 gy, VIB, Leuven, B-3000, Belgium; (3) Cutaneous Biology Research Center, Massachusetts General Hospi14 tal, Harvard Medical School, Charlestown, Massachusetts 02129, USA; (4) Department of Molecular Cell Bi15 ology, Sanquin Research and Landsteiner Laboratory, Academic Medical Centre, University of Amsterdam, 16 1066CX Amsterdam, the Netherlands; (5) Department of Chemistry, University College London, London 17 WC1E6BT, UK; (6) Molecular Imaging and Photonics, University of Leuven, Leuven, B-3000, Belgium; (7) 18 Tytgat Institute for Liver and Gastrointestinal Research, Academic Medical Center University of Amsterdam, 19 1105BK Amsterdam, The Netherlands; (8) Center for Cell Analyses and Modelling, University of Connecticut 20 Health Centre, Farmington 06032, USA; (9) Inflammation Research Center, VIB, Ghent, B-9000, Belgium; 21 (10) Department of Biomedical Molecular Biology, Ghent University, Ghent, B-9000, Belgium; (11) De22 partment of Microbiology and Immunology, University of Leuven, Leuven, B-3000, Belgium; (12) Institute of 23 Structural Molecular Biology, University College London, London WC1E6BT, UK. Present affiliations: (&) 24 Université de Lille, INSERM U1003, Physiologie Cellulaire, F-59000 Lille, France; ($) Max-Delbrück-Center 25 for Molecular Medicine, 13092 Berlin, Germany ; (†) Immunology Department, St. Jude Children’s Research 26 Hospital, Memphis 38105, USA; (‡) DFG-Research Center for Regenerative Therapies, Technical University 27 Dresden, 01307 Dresden, Germany; (#) Department of Biomedical Physiology and Kinesiology, Simon Fra28 ser University, Burnaby V5A 1S6, Canada; (¶) Division of Cardiovascular and Diabetes Research, Multidis29 ciplinary Cardiovascular Research Centre, University of Leeds, Leeds LS29JT, U.K. *equal contribution 30