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Rushabh Gujarathi Section of Hematology and Oncology, Department of Medicine, University of Chicago, Chicago, Illinois, USA

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Sara Abou Azar Section of Endocrine Surgery, Department of Surgery, University of Chicago, Chicago, Illinois, USA

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Joseph Tobias Section of Endocrine Surgery, Department of Surgery, University of Chicago, Chicago, Illinois, USA

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Blase N Polite Section of Hematology and Oncology, Department of Medicine, University of Chicago, Chicago, Illinois, USA

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Namrata Setia Department of Pathology, University of Chicago, Chicago, Illinois, USA

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Nicholas Feinberg Department of Radiology, University of Chicago, Chicago, Illinois, USA

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Daniel E Appelbaum Department of Radiology, University of Chicago, Chicago, Illinois, USA

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Xavier M Keutgen Section of Endocrine Surgery, Department of Surgery, University of Chicago, Chicago, Illinois, USA

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Chih-Yi Liao Section of Hematology and Oncology, Department of Medicine, University of Chicago, Chicago, Illinois, USA

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et al. 2022 ). Mutations in the MEN1 , DAXX (death domain associated protein), and ATRX (ATRX chromatin remodeler) genes are commonly found in NETs, especially in PNETs ( Jiao et al. 2011 , Puccini et al. 2020 ). MEN1 (menin 1) codes

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Christodoulos P Pipinikas Medical Genomics Laboratory, University College London Cancer Institute, London, UK

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Alison M Berner Medical Genomics Laboratory, University College London Cancer Institute, London, UK

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Teresa Sposito Medical Genomics Laboratory, University College London Cancer Institute, London, UK

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Christina Thirlwell Medical Genomics Laboratory, University College London Cancer Institute, London, UK
Neuroendocrine Tumour Unit and Department of Oncology, Royal Free Hospital, London, UK

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disease pathogenesis. Approximately 43% of tumours were found to carry mutations in DAXX (25%) or ATRX (17.6%). Within the cohort, 23.5% of tumours were found to harbour MEN1 mutations in the presence of either DAXX or ATRX , while 20.6% tumours

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Tiago Bordeira Gaspar i3S – Instituto de Investigação e Inovação em Saúde, Porto, Portugal
Ipatimup – Instituto de Patologia e Imunologia Molecular da Universidade do Porto, Porto, Portugal
ICBAS – Instituto de Ciências Biomédicas Abel Salazar da Universidade do Porto, Porto, Portugal
FMUP – Faculdade de Medicina da Universidade do Porto, Porto, Portugal

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José Manuel Lopes i3S – Instituto de Investigação e Inovação em Saúde, Porto, Portugal
Ipatimup – Instituto de Patologia e Imunologia Molecular da Universidade do Porto, Porto, Portugal
FMUP – Faculdade de Medicina da Universidade do Porto, Porto, Portugal
Department of Pathology, Centro Hospitalar e Universitário de São João, Porto, Portugal

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Paula Soares i3S – Instituto de Investigação e Inovação em Saúde, Porto, Portugal
Ipatimup – Instituto de Patologia e Imunologia Molecular da Universidade do Porto, Porto, Portugal
FMUP – Faculdade de Medicina da Universidade do Porto, Porto, Portugal

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João Vinagre i3S – Instituto de Investigação e Inovação em Saúde, Porto, Portugal
Ipatimup – Instituto de Patologia e Imunologia Molecular da Universidade do Porto, Porto, Portugal
FMUP – Faculdade de Medicina da Universidade do Porto, Porto, Portugal

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et al. 2019 , Hong et al. 2020 , Song et al. 2021 ). Besides the menin 1 gene ( MEN1 ) (mutated in 21–44%), novel genes started to be regarded as essential players in PanNEN tumorigenesis, such as the ATRX chromatin remodeler ( ATRX ) (11

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Wenzel M Hackeng Department of Pathology, University Medical Center Utrecht, Utrecht, The Netherlands

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Lodewijk A A Brosens Department of Pathology, University Medical Center Utrecht, Utrecht, The Netherlands

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Koen M A Dreijerink Department of Pathology, University Medical Center Utrecht, Utrecht, The Netherlands
Department of Endocrinology, Amsterdam University Medical Centers, VU University Medical Center, Amsterdam, The Netherlands

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insulinoma. Indolent Aggressive Cases (%) Cases (%) YY1 92/396 (23%) 6/35 (17%) MEN1 10/198 (5%) 1/9 (11%) ALT-phenotype 3/41 (7%) 5/7 (71%) ATRX or DAXX 3/212 (1%) 5/10 (50%) CDKN2A

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M Cives Department of Gastrointestinal Oncology, H. Lee Moffitt Cancer Center and Research Institute

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M Ghayouri Department of Gastrointestinal Oncology, H. Lee Moffitt Cancer Center and Research Institute

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B Morse Department of Gastrointestinal Oncology, H. Lee Moffitt Cancer Center and Research Institute

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M Brelsford Department of Gastrointestinal Oncology, H. Lee Moffitt Cancer Center and Research Institute

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M Black Department of Gastrointestinal Oncology, H. Lee Moffitt Cancer Center and Research Institute

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A Rizzo Department of Pathology, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA

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A Meeker Department of Pathology, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA

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J Strosberg Department of Gastrointestinal Oncology, H. Lee Moffitt Cancer Center and Research Institute

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the correlation between proliferative activity and tumor response. Clinically aggressive pNETs are also characterized by chromosomal instability (CIN) ( Jonkers et al . 2005 ), which has been recently associated with loss of DAXX/ATRX and activation

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I Marinoni Institute of Pathology, University of Bern, Bern, Switzerland

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A Wiederkeher Institute of Pathology, University of Bern, Bern, Switzerland

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T Wiedmer Institute of Pathology, University of Bern, Bern, Switzerland
GCB Graduate School Bern, Bern, Switzerland

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S Pantasis Institute of Pathology, University of Bern, Bern, Switzerland

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A Di Domenico Institute of Pathology, University of Bern, Bern, Switzerland
GCB Graduate School Bern, Bern, Switzerland

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R Frank Institute of Pathology, University of Bern, Bern, Switzerland

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E Vassella Institute of Pathology, University of Bern, Bern, Switzerland

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A Schmitt Institute of Pathology, University of Bern, Bern, Switzerland

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A Perren Institute of Pathology, University of Bern, Bern, Switzerland

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Introduction The molecular pathways and mechanisms underlying initiation and progression of pancreatic neuroendocrine tumors (PanNET) are still poorly understood. Mutations in DAXX (death domain-associated protein) and ATRX (ATR-X) with

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T Vandamme Center of Oncological Research (CORE), University of Antwerp, Antwerp, Belgium
Section of Endocrinology, Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands

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M Beyens Center of Oncological Research (CORE), University of Antwerp, Antwerp, Belgium

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G Boons Center of Oncological Research (CORE), University of Antwerp, Antwerp, Belgium

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A Schepers Center of Medical Genetics, University of Antwerp, Antwerp, Belgium

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K Kamp Section of Endocrinology, Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands

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K Biermann Department of Pathology, Erasmus Medical Center, Rotterdam, The Netherlands

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P Pauwels Department of Pathology, University of Antwerp, Antwerp, Belgium

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W W De Herder Section of Endocrinology, Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands

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L J Hofland Section of Endocrinology, Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands

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M Peeters Center of Oncological Research (CORE), University of Antwerp, Antwerp, Belgium

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G Van Camp Center of Medical Genetics, University of Antwerp, Antwerp, Belgium

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K Op de Beeck Center of Oncological Research (CORE), University of Antwerp, Antwerp, Belgium

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sequenced tumors. Additionally, DAXX was found to be mutated in 22–25% of all tumor samples, while ATRX was mutated in 10–17% of all tumors. Menin (the MEN1 protein), DAXX and ATRX are epigenetic regulators. Menin is involved in histone modification

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Darren Cowzer Department of Medicine, Memorial Sloan Kettering Cancer Center, New York, New York, USA

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Ronak H Shah Kravis Center for Molecular Oncology, Memorial Sloan Kettering Cancer Center, New York, New York, USA

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Joanne F Chou Department of Epidemiology and Biostatistics, Memorial Sloan Kettering Cancer Center, New York, New York, USA

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Ritika Kundra Kravis Center for Molecular Oncology, Memorial Sloan Kettering Cancer Center, New York, New York, USA

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Sippy Punn Department of Medicine, Memorial Sloan Kettering Cancer Center, New York, New York, USA

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Laura Fiedler Department of Medicine, Memorial Sloan Kettering Cancer Center, New York, New York, USA

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April DeMore Department of Medicine, Memorial Sloan Kettering Cancer Center, New York, New York, USA

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Marinela Capanu Department of Epidemiology and Biostatistics, Memorial Sloan Kettering Cancer Center, New York, New York, USA

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Michael F Berger Kravis Center for Molecular Oncology, Memorial Sloan Kettering Cancer Center, New York, New York, USA
Department of Pathology and laboratory medicine, Memorial Sloan Kettering Cancer Center, New York, New York, USA

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Diane Reidy-Lagunes Department of Medicine, Memorial Sloan Kettering Cancer Center, New York, New York, USA
Weill Medical College of Cornell University, New York, New York, USA

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Nitya Raj Department of Medicine, Memorial Sloan Kettering Cancer Center, New York, New York, USA
Weill Medical College of Cornell University, New York, New York, USA

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-differentiated panNENs more frequently harbor alterations in the tumor suppressor gene MEN1 , the chromatin remodeling genes ( DAXX / ATRX ), and in the mTOR pathway ( Jiao et al. 2011 ). In contrast, genetic profiling of poorly differentiated neuroendocrine

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Annunziata Di Domenico Institute of Pathology, University of Bern, Bern, Switzerland
Graduate School for Cellular and Biomedical Sciences, University of Bern, Bern, Switzerland

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Tabea Wiedmer Institute of Pathology, University of Bern, Bern, Switzerland
Graduate School for Cellular and Biomedical Sciences, University of Bern, Bern, Switzerland

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Ilaria Marinoni Institute of Pathology, University of Bern, Bern, Switzerland

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Aurel Perren Institute of Pathology, University of Bern, Bern, Switzerland

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chromatin remodelling and gene expression ( Agarwal et al . 1999 ). The first WES in PanNET reported, in 43% of the cases, mutations in DAXX (death-domain-associated protein) or ATRX (alpha thalassemia/mental retardation syndrome X-linked) ( Jiao et

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A Karpathakis University College London Cancer Institute, Neuroendocrine Tumour Unit, 72 Huntley Street, London WC1E 6BT, UK
University College London Cancer Institute, Neuroendocrine Tumour Unit, 72 Huntley Street, London WC1E 6BT, UK

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H Dibra University College London Cancer Institute, Neuroendocrine Tumour Unit, 72 Huntley Street, London WC1E 6BT, UK

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C Thirlwell University College London Cancer Institute, Neuroendocrine Tumour Unit, 72 Huntley Street, London WC1E 6BT, UK
University College London Cancer Institute, Neuroendocrine Tumour Unit, 72 Huntley Street, London WC1E 6BT, UK

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discovery of mutations in the novel tumour suppressor genes and chromatin remodellers ATRX/DAXX through exome sequencing represent significant progress in our understanding of neuroendocrine tumour (NET) development. NETs are a heterogenous group of

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