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Alfredo Berruti
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Luigi Dogliotti
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, Figueiredo BC, DeLacerda L, Sandrini F, Pianovsky MD, Sampaio G & Sandrini R 2000 Adrenocortical tumors in children. Brazilian Journal of Medical and Biological Research 33 1225 –1234. Schlumberger M , Brugieres L, Gicquel C

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Yunhui Cheng Department of Biological Chemistry, University of Michigan, Ann Arbor, MI, USA

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Raili Emilia Kerppola Atterocor Inc., Ann Arbor, MI, USA

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Tom Klaus Kerppola Department of Biological Chemistry, University of Michigan, Ann Arbor, MI, USA

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caused by ATR-101 in the adrenal cortex of guinea pigs and its apoptotic effects on ACC-derived cells in culture and in xenografts, we speculate that this compound can disrupt mitochondrial functions in tumors of adrenocortical origin. The close

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Anand Pathak Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Douglas R Stewart Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Fabio R Faucz Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Paraskevi Xekouki Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Sara Bass Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Aurelie Vogt Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Xijun Zhang Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Joseph Boland Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Meredith Yeager Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Jennifer T Loud Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Katherine L Nathanson Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Katherine A McGlynn Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Constantine A Stratakis Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Mark H Greene Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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Lisa Mirabello Clinical Genetics Branch, Program on Developmental Endocrinology and Genetics, Cancer Genomics Research Laboratory, Perelman School of Medicine, Genetic Epidemiology Branch, Hormonal and Reproductive Epidemiology Branch, Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, USA

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cyclic AMP signaling in steroidogenic tissue such as the testis, and germline mutations in this gene have been associated with familial adrenocortical tumors ( Horvath et al . 2006 a , b , Kelly 2014 ). Multiple lines of evidence suggest that PDE11A

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Stephanie Espiard Section on Endocrinology & Genetics (SEGEN), Eunice Kennedy Shriver National Institute of Child Health & Human Development (NICHD), National Institutes of Health (NIH), Bethesda, Maryland, USA
Université de Paris, Cochin Institute, Inserm U1016, CNRS 8104, Paris, France

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Ludivine Drougat Section on Endocrinology & Genetics (SEGEN), Eunice Kennedy Shriver National Institute of Child Health & Human Development (NICHD), National Institutes of Health (NIH), Bethesda, Maryland, USA

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Nikolaos Settas Section on Endocrinology & Genetics (SEGEN), Eunice Kennedy Shriver National Institute of Child Health & Human Development (NICHD), National Institutes of Health (NIH), Bethesda, Maryland, USA

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Sara Haydar Section on Endocrinology & Genetics (SEGEN), Eunice Kennedy Shriver National Institute of Child Health & Human Development (NICHD), National Institutes of Health (NIH), Bethesda, Maryland, USA

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Kerstin Bathon Institute of Pharmacology and Toxicology, University of Würzburg, Würzburg, Germany

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Edra London Section on Endocrinology & Genetics (SEGEN), Eunice Kennedy Shriver National Institute of Child Health & Human Development (NICHD), National Institutes of Health (NIH), Bethesda, Maryland, USA

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Isaac Levy Section on Endocrinology & Genetics (SEGEN), Eunice Kennedy Shriver National Institute of Child Health & Human Development (NICHD), National Institutes of Health (NIH), Bethesda, Maryland, USA

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Fabio R Faucz Section on Endocrinology & Genetics (SEGEN), Eunice Kennedy Shriver National Institute of Child Health & Human Development (NICHD), National Institutes of Health (NIH), Bethesda, Maryland, USA

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Davide Calebiro Institute of Metabolism and Systems Research, University of Birmingham, Birmingham, UK
Centre of Membrane Proteins and Receptors (COMPARE), Universities of Nottingham and Birmingham, Birmingham, UK

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Jérôme Bertherat Université de Paris, Cochin Institute, Inserm U1016, CNRS 8104, Paris, France
Center for Rare Adrenal Diseases, Endocrinology Department, Cochin Hospital, Assistance Publique Hôpitaux de Paris, Paris, France

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Dong Li Center for Applied Genomics at The Children’s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA

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Michael A Levine Division of Endocrinology and Diabetes and The Center for Bone Health at The Children’s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA
University of Pennsylvania Perelman School of Medicine, Philadelphia, Pennsylvania, USA

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Constantine A Stratakis Section on Endocrinology & Genetics (SEGEN), Eunice Kennedy Shriver National Institute of Child Health & Human Development (NICHD), National Institutes of Health (NIH), Bethesda, Maryland, USA

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that include the PRKAR1A gene also were described with a variety of skeletal abnormalities ( Salpea et al. 2014 ). PRKACA defects were identified in isolated PPNAD, CPAs and other adrenocortical tumors ( Beuschlein et al. 2014 , Di Dalmazi et

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Y de Keyzer
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D Vieau
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A Picon
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X Bertagna
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A Falchetti Endocrine Unit, Department of Clinical Physiopathology, University of Florence, Florence, Italy

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M L Brandi Endocrine Unit, Department of Clinical Physiopathology, University of Florence, Florence, Italy

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Multiple Endocrine Neoplasias type 1 (MEN 1) and type 2 (MEN 2) represent complex inherited (autosomal dominant traits) syndromes characterized by occurrence of distinct proliferative disorders of endocrine tissues, varying from hyperplasia to adenoma and carcinoma.

MEN 1 syndrome is characterized by parathyroid gland, anterior pituitary and endocrine pancreas tumors. Other endocrine and non endocrine tumors, such as carcinoids, lipomas, pinealomas, adrenocortical and thyroid follicular tumors, have been also described in MEN 1 patients occurring at higher frequency than in general population (Brandi ML et al. 1987). Recently also a spinal ependymoma has been found in a patient with MEN 1 syndrome (Kato H et al 1997)

MEN 2 syndromes recognize three main clinical entities, MEN 2A, characterized by medullary thyroid carcinoma (MTC), primary hyperparathyroidism (PHPT) and pheochromocytoma (PHEO); MEN 2B that exhibits MTC, usually developing sooner than the MEN 2A- associated one, pheochromocytoma, multiple neuromas of gastroenteric mucosa, myelinated corneal nerves (Gorlin RJ et al. 1968) and a typical marphanoid habitus; and familial medullary thyroid carcinoma only (FMTC) featuring by families with at least four members with MTC and no objective evidence of pheochromocytoma and parathyroid disease on screening of affected and at-risk members, as stated by the International RET Mutation Consortium (Larsson C et al. 1994).

Acknowledgements

This work was supported by grants of the Associazione Italiana per la Ricerca sul Cancro (to MLB), from CNR/PF ACRO (INV. 95.00316 PF 39) and by MURST 60% (to MLB).

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I Bossis Section on Endocrinology and Genetics, Developmental Endocrinology Branch, National Institute of Child Health and Human Development, National Institutes of Health, Bethesda, Maryland 20892-1862, USA.

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A Voutetakis Section on Endocrinology and Genetics, Developmental Endocrinology Branch, National Institute of Child Health and Human Development, National Institutes of Health, Bethesda, Maryland 20892-1862, USA.

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T Bei Section on Endocrinology and Genetics, Developmental Endocrinology Branch, National Institute of Child Health and Human Development, National Institutes of Health, Bethesda, Maryland 20892-1862, USA.

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F Sandrini Section on Endocrinology and Genetics, Developmental Endocrinology Branch, National Institute of Child Health and Human Development, National Institutes of Health, Bethesda, Maryland 20892-1862, USA.

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K J Griffin Section on Endocrinology and Genetics, Developmental Endocrinology Branch, National Institute of Child Health and Human Development, National Institutes of Health, Bethesda, Maryland 20892-1862, USA.

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C A Stratakis Section on Endocrinology and Genetics, Developmental Endocrinology Branch, National Institute of Child Health and Human Development, National Institutes of Health, Bethesda, Maryland 20892-1862, USA.

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The type 1 alpha regulatory subunit (R1alpha) of cAMP-dependent protein kinase A (PKA) (PRKAR1A) is an important regulator of the serine-threonine kinase activity catalyzed by the PKA holoenzyme. Carney complex (CNC) describes the association 'of spotty skin pigmentation, myxomas, and endocrine overactivity'; CNC is in essence the latest form of multiple endocrine neoplasia to be described and affects the pituitary, thyroid, adrenal and gonadal glands. Primary pigmented nodular adrenocortical disease (PPNAD), a micronodular form of bilateral adrenal hyperplasia that causes a unique, inherited form of Cushing syndrome, is also the most common endocrine manifestation of CNC. CNC and PPNAD are genetically heterogeneous but one of the responsible genes is PRKAR1A, at least for those families that map to 17q22-24 (the chromosomal region that harbors PRKAR1A). CNC and/or PPNAD are the first human diseases to be caused by mutations in one of the subunits of the PKA holoenzyme. Despite the extensive literature on R1alpha and PKA, little is known about their potential involvement in cell cycle regulation, growth and/or proliferation. The presence of inactivating germline mutations and the loss of its wild-type allele in CNC lesions indicated that PRKAR1A could function as a tumor-suppressor gene in these tissues. However, there are conflicting data in the literature about PRKAR1A's role in human neoplasms, cancer cell lines and animal models. In this report, we review briefly the genetics of CNC and focus on the involvement of PRKAR1A in human tumorigenesis in an effort to reconcile the often diametrically opposite reports on R1alpha.

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S G Creemers Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands
Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands

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L J Hofland Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands
Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands

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E Korpershoek Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands
Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands

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G J H Franssen Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands
Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands

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F J van Kemenade Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands
Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands

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W W de Herder Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands
Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands

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R A Feelders Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands
Rotterdam Adrenal Center, Department of Internal Medicine, Department of Pathology, Department of Surgery, Erasmus MC University Medical Center, Rotterdam, The Netherlands

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during surgery ( Song et al . 2011 ). Pathology The Weiss score (WS) is currently the most widely used classification system for the pathological assessment of adrenocortical tumors ( Weiss 1984 , Lau & Weiss 2009 ). It consists of nine morphological

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Maria Riedmeier University Children’s Hospital, Department of Pediatric Hematology, Oncology and Stem cell transplantation, University of Wuerzburg, Wuerzburg, Germany.

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Lester D R Thompson Head and Neck Pathology Consultations, Los Angeles, California, USA

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Carlos Augusto Fernandes Molina Department of Surgery and Anatomy, Ribeirao Preto Medical School, University of Sao Paulo, Ribeirao Preto, Sao Paulo, Brazil

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Boris Decarolis Department of Pediatric Oncology and Hematology, University Children’s Hospital of Cologne, Medical Faculty, Cologne, Germany

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Christoph Härtel University Children’s Hospital, Department of Pediatric Hematology, Oncology and Stem cell transplantation, University of Wuerzburg, Wuerzburg, Germany.

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Paul-G Schlegel University Children’s Hospital, Department of Pediatric Hematology, Oncology and Stem cell transplantation, University of Wuerzburg, Wuerzburg, Germany.
Comprehensive Cancer Centre Mainfranken, University of Wuerzburg Medical Centre, Wuerzburg, Germany

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Martin Fassnacht Comprehensive Cancer Centre Mainfranken, University of Wuerzburg Medical Centre, Wuerzburg, Germany
Department of Medicine, Division of Endocrinology and Diabetes, University Hospital, University of Wuerzburg, Wuerzburg, Germany

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Verena Wiegering University Children’s Hospital, Department of Pediatric Hematology, Oncology and Stem cell transplantation, University of Wuerzburg, Wuerzburg, Germany.
Comprehensive Cancer Centre Mainfranken, University of Wuerzburg Medical Centre, Wuerzburg, Germany

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hormone profile and seem to have a more favorable outcome ( Mendonca et al. 1995 , Riedmeier et al. 2021 , Zambaiti et al. 2021 ). Several authors have discussed the hypothesis of adrenocortical tumor development in the fetal zone of the adrenal

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Uriel Clemente-Gutierrez Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Carolina R C Pieterman Department of Endocrine Oncology, University Medical Center Utrecht, Utrecht, The Netherlands

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Michael S Lui Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Thomas Szabo Yamashita Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Andrés Tame-Elorduy Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Bernice L Huang Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Aditya S Shirali Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Derek J Erstad Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Jeffrey E Lee Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Sarah B Fisher Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Paul H Graham Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Elizabeth G Grubbs Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Steven G Waguespack Department of Endocrine Neoplasia and Hormonal Disorders, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Chaan S Ng Department of Diagnostic Radiology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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Nancy Perrier Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, Texas, USA

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, information about the natural history of adrenal lesions in MEN1 is scarce. Most MEN1-related adrenal tumors reported in the literature are nonfunctional, benign adrenocortical tumors, although functional cortical tumors, including aldosteronomas ( Beckers

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