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I Savchuk Department of Women’s and Children’s Health, Pediatric Endocrinology Unit, Karolinska Institute & University Hospital, Stockholm, Sweden

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M L Morvan LUNAM Université, École Nationale Vétérinaire, Agroalimentaire et de l’Alimentation, Nantes-Atlantique (Oniris), Laboratoire d’Étude des Résidus et Contaminants dans les Aliments (LABERCA), USC INRA 1329, Nantes, France

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J P Antignac LUNAM Université, École Nationale Vétérinaire, Agroalimentaire et de l’Alimentation, Nantes-Atlantique (Oniris), Laboratoire d’Étude des Résidus et Contaminants dans les Aliments (LABERCA), USC INRA 1329, Nantes, France

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K Gemzell-Danielsson Department of Obstetrics and Gynecology, Karolinska Institute & University Hospital, Stockholm, Sweden

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B Le Bizec LUNAM Université, École Nationale Vétérinaire, Agroalimentaire et de l’Alimentation, Nantes-Atlantique (Oniris), Laboratoire d’Étude des Résidus et Contaminants dans les Aliments (LABERCA), USC INRA 1329, Nantes, France

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O Söder Department of Women’s and Children’s Health, Pediatric Endocrinology Unit, Karolinska Institute & University Hospital, Stockholm, Sweden

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K Svechnikov Department of Women’s and Children’s Health, Pediatric Endocrinology Unit, Karolinska Institute & University Hospital, Stockholm, Sweden

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Introduction Steroid hormones produced by the human fetal adrenal glands (HFA) have been proposed to regulate intrauterine homeostasis and the maturation of certain organs required for extrauterine life ( 1 ). In this case, appropriate

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Manon Engels Department of Paediatrics, Radboud Amalia Children’s Hospital, Radboud university medical center, Nijmegen, The Netherlands
Laboratory Medicine, Radboud Institute for Molecular Life Sciences (RIMLS), Radboud university medical center, Nijmegen, The Netherlands

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Paul N Span Radiation Oncology, Radiotherapy and OncoImmunology Laboratory, RIMLS, Radboud university medical center, Nijmegen, The Netherlands

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Rod T Mitchell MRC Centre for Reproductive Health, University of Edinburgh, The Queen’s Medical Research Institute, Edinburgh, UK

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Joop J T M Heuvel Laboratory Medicine, Radboud Institute for Molecular Life Sciences (RIMLS), Radboud university medical center, Nijmegen, The Netherlands

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Monica A Marijnissen-van Zanten Department of Pathology, Radboud university medical center, Nijmegen, The Netherlands

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Antonius E van Herwaarden Laboratory Medicine, Radboud Institute for Molecular Life Sciences (RIMLS), Radboud university medical center, Nijmegen, The Netherlands

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Christina A Hulsbergen-van de Kaa Department of Pathology, Radboud university medical center, Nijmegen, The Netherlands

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Egbert Oosterwijk Department of Urology, Radboud university medical center, Nijmegen, The Netherlands

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Nike M Stikkelbroeck Department of Internal Medicine, Radboud university medical center, Nijmegen, The Netherlands

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Lee B Smith MRC Centre for Reproductive Health, University of Edinburgh, The Queen’s Medical Research Institute, Edinburgh, UK

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Fred C G J Sweep Laboratory Medicine, Radboud Institute for Molecular Life Sciences (RIMLS), Radboud university medical center, Nijmegen, The Netherlands

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Hedi L Claahsen-van der Grinten Department of Paediatrics, Radboud Amalia Children’s Hospital, Radboud university medical center, Nijmegen, The Netherlands

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.1210/en.2003-0472 12959982 39 Kiiveri S Liu J Westerholm-Ormio M Narita N Wilson DB Voutilainen R Heikinheimo M. Differential expression of GATA-4 and GATA-6 in fetal and adult mouse and human adrenal tissue . Endocrinology

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Deirdre Green Academic Department of Endocrinology, Beaumont Hospital and the Royal College of Surgeons in Ireland, Dublin

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Rosemary Dineen Academic Department of Endocrinology, Beaumont Hospital and the Royal College of Surgeons in Ireland, Dublin

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Michael W O’Reilly Academic Department of Endocrinology, Beaumont Hospital and the Royal College of Surgeons in Ireland, Dublin

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Mark Sherlock Academic Department of Endocrinology, Beaumont Hospital and the Royal College of Surgeons in Ireland, Dublin

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positive feedback loop. Placental CRH stimulates production of fetal cortisol, both by increasing fetal adrenal responsiveness to ACTH and by direct stimulation of the adrenal. Increased fetal cortisol would in turn stimulate placental CRH production

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Stine Linding Andersen Department of Clinical Biochemistry, Aalborg University Hospital, Aalborg, Denmark
Department of Clinical Medicine, Aalborg University, Aalborg, Denmark

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Stig Andersen Department of Clinical Medicine, Aalborg University, Aalborg, Denmark
Department of Geriatrics, Aalborg University Hospital, Aalborg, Denmark

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hyperthyroidism and supporting evidence from experimental and human studies, and we discuss methodological aspects and implications for scientific and clinical practice. The hypothesis of fetal programming Fetal programming is a concept within reproductive

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Fiona Broughton Pipkin Department of Obstetrics and Gynaecology, Department of Nephrology, Leicester Royal Infirmary, School of Medicine, University of Nottingham, Nottingham, NG5 1PB, UK

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Hiten D Mistry Department of Obstetrics and Gynaecology, Department of Nephrology, Leicester Royal Infirmary, School of Medicine, University of Nottingham, Nottingham, NG5 1PB, UK

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Chandrima Roy Department of Obstetrics and Gynaecology, Department of Nephrology, Leicester Royal Infirmary, School of Medicine, University of Nottingham, Nottingham, NG5 1PB, UK

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Bernhard Dick Department of Obstetrics and Gynaecology, Department of Nephrology, Leicester Royal Infirmary, School of Medicine, University of Nottingham, Nottingham, NG5 1PB, UK

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Jason Waugh Department of Obstetrics and Gynaecology, Department of Nephrology, Leicester Royal Infirmary, School of Medicine, University of Nottingham, Nottingham, NG5 1PB, UK

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Rebecca Chikhi Department of Obstetrics and Gynaecology, Department of Nephrology, Leicester Royal Infirmary, School of Medicine, University of Nottingham, Nottingham, NG5 1PB, UK

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Lesia O Kurlak Department of Obstetrics and Gynaecology, Department of Nephrology, Leicester Royal Infirmary, School of Medicine, University of Nottingham, Nottingham, NG5 1PB, UK

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Markus G Mohaupt Department of Obstetrics and Gynaecology, Department of Nephrology, Leicester Royal Infirmary, School of Medicine, University of Nottingham, Nottingham, NG5 1PB, UK

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153 – 165 . ( doi:10.1530/eje.0.1350153 ). 2 Rainey WE Rehman KS Carr BR . Fetal and maternal adrenals in human pregnancy . Obstetrics and Gynecology Clinics of North America 2004 31 817 – 835 . ( doi:10.1016/j.ogc.2004.08.006 ). 3

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Erika Peverelli Endocrinology and Diabetology Unit, Diabetology and Metabolic Disease Unit, Endocrinology Unit, Unit of Human Pathology, Surgical Department, Department of Clinical Sciences and Community Health, University of Milan, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Pad.Granelli, Via F. Sforza, 35, 20122 Milan, Italy

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Federica Ermetici Endocrinology and Diabetology Unit, Diabetology and Metabolic Disease Unit, Endocrinology Unit, Unit of Human Pathology, Surgical Department, Department of Clinical Sciences and Community Health, University of Milan, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Pad.Granelli, Via F. Sforza, 35, 20122 Milan, Italy
Endocrinology and Diabetology Unit, Diabetology and Metabolic Disease Unit, Endocrinology Unit, Unit of Human Pathology, Surgical Department, Department of Clinical Sciences and Community Health, University of Milan, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Pad.Granelli, Via F. Sforza, 35, 20122 Milan, Italy

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Sabrina Corbetta Endocrinology and Diabetology Unit, Diabetology and Metabolic Disease Unit, Endocrinology Unit, Unit of Human Pathology, Surgical Department, Department of Clinical Sciences and Community Health, University of Milan, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Pad.Granelli, Via F. Sforza, 35, 20122 Milan, Italy

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Ettore Gozzini Endocrinology and Diabetology Unit, Diabetology and Metabolic Disease Unit, Endocrinology Unit, Unit of Human Pathology, Surgical Department, Department of Clinical Sciences and Community Health, University of Milan, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Pad.Granelli, Via F. Sforza, 35, 20122 Milan, Italy

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Laura Avagliano Endocrinology and Diabetology Unit, Diabetology and Metabolic Disease Unit, Endocrinology Unit, Unit of Human Pathology, Surgical Department, Department of Clinical Sciences and Community Health, University of Milan, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Pad.Granelli, Via F. Sforza, 35, 20122 Milan, Italy

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Marco A Zappa Endocrinology and Diabetology Unit, Diabetology and Metabolic Disease Unit, Endocrinology Unit, Unit of Human Pathology, Surgical Department, Department of Clinical Sciences and Community Health, University of Milan, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Pad.Granelli, Via F. Sforza, 35, 20122 Milan, Italy

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Gaetano Bulfamante Endocrinology and Diabetology Unit, Diabetology and Metabolic Disease Unit, Endocrinology Unit, Unit of Human Pathology, Surgical Department, Department of Clinical Sciences and Community Health, University of Milan, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Pad.Granelli, Via F. Sforza, 35, 20122 Milan, Italy

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Paolo Beck-Peccoz Endocrinology and Diabetology Unit, Diabetology and Metabolic Disease Unit, Endocrinology Unit, Unit of Human Pathology, Surgical Department, Department of Clinical Sciences and Community Health, University of Milan, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Pad.Granelli, Via F. Sforza, 35, 20122 Milan, Italy

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Anna Spada Endocrinology and Diabetology Unit, Diabetology and Metabolic Disease Unit, Endocrinology Unit, Unit of Human Pathology, Surgical Department, Department of Clinical Sciences and Community Health, University of Milan, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Pad.Granelli, Via F. Sforza, 35, 20122 Milan, Italy

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Giovanna Mantovani Endocrinology and Diabetology Unit, Diabetology and Metabolic Disease Unit, Endocrinology Unit, Unit of Human Pathology, Surgical Department, Department of Clinical Sciences and Community Health, University of Milan, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Pad.Granelli, Via F. Sforza, 35, 20122 Milan, Italy

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are still scarce. In this study, we evaluated the expression of PKA regulatory subunits in human adipose tissue during fetal development as well as their role in the differentiation process of two in vitro cell models of adipogenesis, i.e., 3T3-L1

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Tatiana V Novoselova Centre for Endocrinology, William Harvey Research Institute, Barts and the London School of Medicine, Queen Mary University of London, London, UK

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Peter J King Centre for Endocrinology, William Harvey Research Institute, Barts and the London School of Medicine, Queen Mary University of London, London, UK

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Leonardo Guasti Centre for Endocrinology, William Harvey Research Institute, Barts and the London School of Medicine, Queen Mary University of London, London, UK

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Louise A Metherell Centre for Endocrinology, William Harvey Research Institute, Barts and the London School of Medicine, Queen Mary University of London, London, UK

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Adrian J L Clark Centre for Endocrinology, William Harvey Research Institute, Barts and the London School of Medicine, Queen Mary University of London, London, UK

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Li F Chan Centre for Endocrinology, William Harvey Research Institute, Barts and the London School of Medicine, Queen Mary University of London, London, UK

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Hypothalamo–pituitary–adrenal axis The hypothalamo–pituitary–adrenal (HPA) axis dictates the production of glucocorticoids secreted from the adrenal gland. Parvocellular neurosecretory neurons within the hypothalamic paraventricular nucleus

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Emanuelle Nunes-Souza Pelé Pequeno Príncipe Research Institute, Água Verde, Curitiba, Parana, Brazil
Faculdades Pequeno Príncipe, Rebouças, Curitiba, Parana, Brazil
Centro de Genética Molecular e Pesquisa do Câncer em Crianças (CEGEMPAC) at Universidade Federal do Paraná, Agostinho Leão Jr., Glória, Curitiba, Parana, Brazil

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Mônica Evelise Silveira Laboratório Central de Análises Clínicas, Hospital de Clínicas, Universidade Federal do Paraná, Centro, Curitiba, Paraná, Brazil

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Monalisa Castilho Mendes Pelé Pequeno Príncipe Research Institute, Água Verde, Curitiba, Parana, Brazil
Faculdades Pequeno Príncipe, Rebouças, Curitiba, Parana, Brazil
Centro de Genética Molecular e Pesquisa do Câncer em Crianças (CEGEMPAC) at Universidade Federal do Paraná, Agostinho Leão Jr., Glória, Curitiba, Parana, Brazil

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Seigo Nagashima Serviço de Anatomia Patológica, Hospital de Clínicas, Universidade Federal do Paraná, General Carneiro, Alto da Glória, Curitiba, Parana, Brazil
Departamento de Medicina, PUC-PR, Prado Velho, Curitiba, Parana, Brazil

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Caroline Busatta Vaz de Paula Serviço de Anatomia Patológica, Hospital de Clínicas, Universidade Federal do Paraná, General Carneiro, Alto da Glória, Curitiba, Parana, Brazil
Departamento de Medicina, PUC-PR, Prado Velho, Curitiba, Parana, Brazil

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Guilherme Guilherme Vieira Cavalcante da Silva Serviço de Anatomia Patológica, Hospital de Clínicas, Universidade Federal do Paraná, General Carneiro, Alto da Glória, Curitiba, Parana, Brazil
Departamento de Medicina, PUC-PR, Prado Velho, Curitiba, Parana, Brazil

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Giovanna Silva Barbosa Serviço de Anatomia Patológica, Hospital de Clínicas, Universidade Federal do Paraná, General Carneiro, Alto da Glória, Curitiba, Parana, Brazil
Departamento de Medicina, PUC-PR, Prado Velho, Curitiba, Parana, Brazil

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Julia Belgrowicz Martins Pelé Pequeno Príncipe Research Institute, Água Verde, Curitiba, Parana, Brazil
Faculdades Pequeno Príncipe, Rebouças, Curitiba, Parana, Brazil

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Lúcia de Noronha Serviço de Anatomia Patológica, Hospital de Clínicas, Universidade Federal do Paraná, General Carneiro, Alto da Glória, Curitiba, Parana, Brazil
Departamento de Medicina, PUC-PR, Prado Velho, Curitiba, Parana, Brazil

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Luana Lenzi Departamento de Análises Clínicas, Universidade Federal do Paraná, Curitiba, Paraná, Brazil

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José Renato Sales Barbosa Pelé Pequeno Príncipe Research Institute, Água Verde, Curitiba, Parana, Brazil
Centro de Genética Molecular e Pesquisa do Câncer em Crianças (CEGEMPAC) at Universidade Federal do Paraná, Agostinho Leão Jr., Glória, Curitiba, Parana, Brazil

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Rayssa Danilow Fachin Donin Centro de Genética Molecular e Pesquisa do Câncer em Crianças (CEGEMPAC) at Universidade Federal do Paraná, Agostinho Leão Jr., Glória, Curitiba, Parana, Brazil

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Juliana Ferreira de Moura Pós Graduação em Microbiologia, Parasitologia e Patologia, Departamento de Patologia Básica – UFPR, Curitiba, Brazil

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Gislaine Custódio Faculdades Pequeno Príncipe, Rebouças, Curitiba, Parana, Brazil
Laboratório Central de Análises Clínicas, Hospital de Clínicas, Universidade Federal do Paraná, Centro, Curitiba, Paraná, Brazil

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Cleber Machado-Souza Pelé Pequeno Príncipe Research Institute, Água Verde, Curitiba, Parana, Brazil
Faculdades Pequeno Príncipe, Rebouças, Curitiba, Parana, Brazil
Centro de Genética Molecular e Pesquisa do Câncer em Crianças (CEGEMPAC) at Universidade Federal do Paraná, Agostinho Leão Jr., Glória, Curitiba, Parana, Brazil

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Enzo Lalli Institut de Pharmacologie Moléculaire et Cellulaire CNRS, Sophia Antipolis, Valbonne, France

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Bonald Cavalcante de Figueiredo Pelé Pequeno Príncipe Research Institute, Água Verde, Curitiba, Parana, Brazil
Faculdades Pequeno Príncipe, Rebouças, Curitiba, Parana, Brazil
Centro de Genética Molecular e Pesquisa do Câncer em Crianças (CEGEMPAC) at Universidade Federal do Paraná, Agostinho Leão Jr., Glória, Curitiba, Parana, Brazil
Departamento de Saúde Coletiva, Universidade Federal do Paraná, Curitiba, Paraná, Brazil

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-008-9105-1 ) 5 Mason JI Rainey WE Steroidogenesis in the human fetal adrenal: a role for cholesterol synthesized de novo . Journal of Clinical Endocrinology and Metabolism 1987 64 140 – 147 . ( https://doi.org/10.1210/jcem-64-1-140 ) 6 Ishimoto H

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Annelies van’t Westeinde Department of Women’s and Children’s Health, Karolinska Institutet and Division of Pediatrics, Unit for Pediatric Endocrinology and Metabolic Disorders, Karolinska University Hospital, Stockholm, Sweden

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Leif Karlsson Department of Women’s and Children’s Health, Karolinska Institutet and Division of Pediatrics, Unit for Pediatric Endocrinology and Metabolic Disorders, Karolinska University Hospital, Stockholm, Sweden

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Valeria Messina Department of Women’s and Children’s Health, Karolinska Institutet and Division of Pediatrics, Unit for Pediatric Endocrinology and Metabolic Disorders, Karolinska University Hospital, Stockholm, Sweden

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Lena Wallensteen Department of Women’s and Children’s Health, Karolinska Institutet and Division of Pediatrics, Unit for Pediatric Endocrinology and Metabolic Disorders, Karolinska University Hospital, Stockholm, Sweden

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Manuela Brösamle European Patient Advocacy Group for Adrenal Diseases, European Reference Network on Rare Endocrine Conditions (Endo ERN), Endo ERN Coordinating Centre, Leiden, The Netherlands

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Giorgio Dal Maso ArfSAG (Associazione Refionale Famiglie Sindrome Adreno Genitale) c/o Unita Operativa di Pediatria, Azienda Ospedaliero Universitaria di Bologna, Policlinico S Orsala-Malpighi, Bologna, Italy

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Alessandro Lazzerini Spanish Association of Congenital Adrenal Hyperplasia (CAH), Spain

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Jette Kristensen ePAG & Chair of Danish Addison Patient Association, Aarhus, Denmark

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Diana Kwast Dutch Adrenal Society NVACP, Nijkerk, The Netherlands

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Lea Tschaidse Department of Endocrinology, Medizinische Klinik IV, Klinikum der Universität München, Munich, Germany

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Matthias K Auer Department of Endocrinology, Medizinische Klinik IV, Klinikum der Universität München, Munich, Germany

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Hanna F Nowotny Department of Endocrinology, Medizinische Klinik IV, Klinikum der Universität München, Munich, Germany

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Luca Persani Department of Medical Biotechnology and Translational Medicine, University of Milan, Milan, Italy
Department of Endocrine and Metabolic Diseases, Istituto Auxologico Italiano IRCCS, Milan, Italy

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Nicole Reisch Department of Endocrinology, Medizinische Klinik IV, Klinikum der Universität München, Munich, Germany

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Svetlana Lajic Department of Women’s and Children’s Health, Karolinska Institutet and Division of Pediatrics, Unit for Pediatric Endocrinology and Metabolic Disorders, Karolinska University Hospital, Stockholm, Sweden

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RB . Development and function of the human fetal adrenal cortex: a key component in the feto-placental unit . Endocrine Reviews 2011 32 317 – 355 . ( https://doi.org/10.1210/er.2010-0001 ) 41 Mesiano S Jaffe RB . Developmental and functional

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Stavroula A Paschou Department of Endocrinology and Diabetes, Hellenic Red Cross Hospital, Athens, Greece

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Eleni Palioura Department of Endocrinology and Diabetes, Hellenic Red Cross Hospital, Athens, Greece

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Dimitrios Ioannidis Department of Endocrinology and Diabetes, Sismanoglio-Amalia Fleming Hospital, Athens, Greece

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Panagiotis Anagnostis Unit of Reproductive Endocrinology, First Department of Obstetrics and Gynecology, Medical School, Aristotle University of Thessaloniki, Thessaloniki, Greece

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Argyro Panagiotakou Department of Endocrinology and Diabetes, Sismanoglio-Amalia Fleming Hospital, Athens, Greece

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Vasiliki Loi Department of Endocrinology and Diabetes, Hellenic Red Cross Hospital, Athens, Greece

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Georgios Karageorgos Department of Endocrinology and Diabetes, Sismanoglio-Amalia Fleming Hospital, Athens, Greece

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Dimitrios G Goulis Unit of Reproductive Endocrinology, First Department of Obstetrics and Gynecology, Medical School, Aristotle University of Thessaloniki, Thessaloniki, Greece

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Andromachi Vryonidou Department of Endocrinology and Diabetes, Hellenic Red Cross Hospital, Athens, Greece

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degree of insulin resistance ( 7 ). Androgens are mainly of ovarian origin but in 30% of the patients, they originate from the adrenal glands ( 8 ), the main representative being dehydroepiandrosterone sulfate (DHEA-S). The effect of this specific (mainly

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