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Johanna L Barclay School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia
School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia
School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia

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Hadiya Agada School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia
School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia

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Christina Jang School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia
School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia

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Micheal Ward School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia
School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia

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Neil Wetzig School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia

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Ken K Y Ho School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia
School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia
School of Medicine, Mater Research Institute, The Translational Research Institute, Princess Alexandra Hospital, University of Queensland, Herston, Queensland, Australia

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activated BAT. These observations provide persuasive evidence for a significant metabolic role of BAT in humans. Glucocorticoids (GCs) play a critical role in the regulation of energy balance. Chronic GC excess, as exemplified in Cushing's syndrome, results

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Francesca Spiga Henry Wellcome Laboratories for Integrative Neuroscience and Endocrinology, College of Engineering, Wellcome Trust Centre for Biomedical Modelling and Analysis, University of Bristol, Dorothy Hodgkin Building, Whitson Street, Bristol BS1 3NY, UK

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Jamie J Walker Henry Wellcome Laboratories for Integrative Neuroscience and Endocrinology, College of Engineering, Wellcome Trust Centre for Biomedical Modelling and Analysis, University of Bristol, Dorothy Hodgkin Building, Whitson Street, Bristol BS1 3NY, UK
Henry Wellcome Laboratories for Integrative Neuroscience and Endocrinology, College of Engineering, Wellcome Trust Centre for Biomedical Modelling and Analysis, University of Bristol, Dorothy Hodgkin Building, Whitson Street, Bristol BS1 3NY, UK
Henry Wellcome Laboratories for Integrative Neuroscience and Endocrinology, College of Engineering, Wellcome Trust Centre for Biomedical Modelling and Analysis, University of Bristol, Dorothy Hodgkin Building, Whitson Street, Bristol BS1 3NY, UK

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Rita Gupta Henry Wellcome Laboratories for Integrative Neuroscience and Endocrinology, College of Engineering, Wellcome Trust Centre for Biomedical Modelling and Analysis, University of Bristol, Dorothy Hodgkin Building, Whitson Street, Bristol BS1 3NY, UK

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John R Terry Henry Wellcome Laboratories for Integrative Neuroscience and Endocrinology, College of Engineering, Wellcome Trust Centre for Biomedical Modelling and Analysis, University of Bristol, Dorothy Hodgkin Building, Whitson Street, Bristol BS1 3NY, UK
Henry Wellcome Laboratories for Integrative Neuroscience and Endocrinology, College of Engineering, Wellcome Trust Centre for Biomedical Modelling and Analysis, University of Bristol, Dorothy Hodgkin Building, Whitson Street, Bristol BS1 3NY, UK

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Stafford L Lightman Henry Wellcome Laboratories for Integrative Neuroscience and Endocrinology, College of Engineering, Wellcome Trust Centre for Biomedical Modelling and Analysis, University of Bristol, Dorothy Hodgkin Building, Whitson Street, Bristol BS1 3NY, UK

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Introduction Glucocorticoids, the end-product of the hypothalamic–pituitary–adrenal (HPA) axis, are essential hormones that regulate the organism's homeostasis and its response to stress. Glucocorticoids (corticosterone in the rat, cortisol in

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Miski Scerif Centre for Endocrinology, Department of Endocrine Neurobiology, Division of Endocrinology, Oxford Centre for Diabetes, William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK

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Tamás Füzesi Centre for Endocrinology, Department of Endocrine Neurobiology, Division of Endocrinology, Oxford Centre for Diabetes, William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK

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Julia D Thomas Centre for Endocrinology, Department of Endocrine Neurobiology, Division of Endocrinology, Oxford Centre for Diabetes, William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK

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Blerina Kola Centre for Endocrinology, Department of Endocrine Neurobiology, Division of Endocrinology, Oxford Centre for Diabetes, William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK

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Ashley B Grossman Centre for Endocrinology, Department of Endocrine Neurobiology, Division of Endocrinology, Oxford Centre for Diabetes, William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK
Centre for Endocrinology, Department of Endocrine Neurobiology, Division of Endocrinology, Oxford Centre for Diabetes, William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK

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Csaba Fekete Centre for Endocrinology, Department of Endocrine Neurobiology, Division of Endocrinology, Oxford Centre for Diabetes, William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK
Centre for Endocrinology, Department of Endocrine Neurobiology, Division of Endocrinology, Oxford Centre for Diabetes, William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK

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Márta Korbonits Centre for Endocrinology, Department of Endocrine Neurobiology, Division of Endocrinology, Oxford Centre for Diabetes, William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK

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Introduction Cushing's syndrome results from chronic exposure to high doses of glucocorticoids. Endogenous Cushing's syndrome is rare and may be caused by an adrenocorticotrophin (ACTH)-secreting pituitary adenoma (Cushing's disease) or a cortisol

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Ying Sze Centre for Discovery Brain Sciences, University of Edinburgh, Edinburgh, UK
The Roslin Institute, University of Edinburgh, Easter Bush Campus, Midlothian, UK

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Joana Fernandes The Roslin Institute, University of Edinburgh, Easter Bush Campus, Midlothian, UK

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Zofia M Kołodziejczyk Centre for Discovery Brain Sciences, University of Edinburgh, Edinburgh, UK

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Paula J Brunton Centre for Discovery Brain Sciences, University of Edinburgh, Edinburgh, UK
The Roslin Institute, University of Edinburgh, Easter Bush Campus, Midlothian, UK
Zhejiang University-University of Edinburgh Institute, International Campus, Haining, Zhejiang, P.R. China

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( Pariante & Lightman 2008 ). The HPA axis is the primary neuroendocrine stress response system. It is activated following perturbations to normal homeostasis, resulting in increased secretion of glucocorticoids from the adrenal gland, which mobilises energy

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Michaela D Wharfe School of Anatomy, Metabolomics Australia, Physiology and Human Biology, The University of Western Australia, M309, Perth 6009, Australia

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Peter J Mark School of Anatomy, Metabolomics Australia, Physiology and Human Biology, The University of Western Australia, M309, Perth 6009, Australia

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Caitlin S Wyrwoll School of Anatomy, Metabolomics Australia, Physiology and Human Biology, The University of Western Australia, M309, Perth 6009, Australia

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Jeremy T Smith School of Anatomy, Metabolomics Australia, Physiology and Human Biology, The University of Western Australia, M309, Perth 6009, Australia

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Cassandra Yap School of Anatomy, Metabolomics Australia, Physiology and Human Biology, The University of Western Australia, M309, Perth 6009, Australia

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Michael W Clarke School of Anatomy, Metabolomics Australia, Physiology and Human Biology, The University of Western Australia, M309, Perth 6009, Australia

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Brendan J Waddell School of Anatomy, Metabolomics Australia, Physiology and Human Biology, The University of Western Australia, M309, Perth 6009, Australia

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–pituitary–adrenal (HPA) axis after mid-gestation leads to increased glucocorticoid levels ( Patrick et al . 1980 , Atkinson & Waddell 1995 ). This HPA axis adaptation is crucial because it promotes the release of energy stores to meet high fetal demand ( Atkinson

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Huali Yu Key Laboratory of Molecular Epigenetics, Ministry of Education and Institute of Cytology and Genetics, Northeast Normal University, Changchun, China

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Ye Guo Key Laboratory of Molecular Epigenetics, Ministry of Education and Institute of Cytology and Genetics, Northeast Normal University, Changchun, China

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Yang Zhao Key Laboratory of Molecular Epigenetics, Ministry of Education and Institute of Cytology and Genetics, Northeast Normal University, Changchun, China

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Feng Zhou Key Laboratory of Molecular Epigenetics, Ministry of Education and Institute of Cytology and Genetics, Northeast Normal University, Changchun, China

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Kehan Zhao Key Laboratory of Molecular Epigenetics, Ministry of Education and Institute of Cytology and Genetics, Northeast Normal University, Changchun, China

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Mayuqing Li Key Laboratory of Molecular Epigenetics, Ministry of Education and Institute of Cytology and Genetics, Northeast Normal University, Changchun, China

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Junxiong Wen Key Laboratory of Molecular Epigenetics, Ministry of Education and Institute of Cytology and Genetics, Northeast Normal University, Changchun, China

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Zixuan He Key Laboratory of Molecular Epigenetics, Ministry of Education and Institute of Cytology and Genetics, Northeast Normal University, Changchun, China

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Xiaojuan Zhu Key Laboratory of Molecular Epigenetics, Ministry of Education and Institute of Cytology and Genetics, Northeast Normal University, Changchun, China

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Xiaoxiao He Key Laboratory of Molecular Epigenetics, Ministry of Education and Institute of Cytology and Genetics, Northeast Normal University, Changchun, China

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Introduction Glucocorticoids (GCs) have profound influences on many physiological functions, including growth, metabolism, development, behaviors and stress reactions ( Blodgett et al. 1956 , Munck et al. 1984 , Haskett 1985 ). They also

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K S Wilson The University/BHF Centre for Cardiovascular Science, University of Edinburgh, The Queen’s Medical Research Institute, Edinburgh, UK

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C S Tucker The University/BHF Centre for Cardiovascular Science, University of Edinburgh, The Queen’s Medical Research Institute, Edinburgh, UK

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E A S Al-Dujaili The University/BHF Centre for Cardiovascular Science, University of Edinburgh, The Queen’s Medical Research Institute, Edinburgh, UK

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M C Holmes The University/BHF Centre for Cardiovascular Science, University of Edinburgh, The Queen’s Medical Research Institute, Edinburgh, UK

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P W F Hadoke The University/BHF Centre for Cardiovascular Science, University of Edinburgh, The Queen’s Medical Research Institute, Edinburgh, UK

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C J Kenyon The University/BHF Centre for Cardiovascular Science, University of Edinburgh, The Queen’s Medical Research Institute, Edinburgh, UK

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M A Denvir The University/BHF Centre for Cardiovascular Science, University of Edinburgh, The Queen’s Medical Research Institute, Edinburgh, UK

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publications have highlighted the developmental aspects of altered embryonic glucocorticoid exposure in zebrafish ( Hillegass et al . 2008 , Pikulkaew et al . 2011 , De Marco et al . 2013 , Nesan & Vijayan 2013 , 2016 , Faught et al . 2016 ), few

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P Nguyen Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada
Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada

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S Khurana Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada

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H Peltsch Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada
Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada

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J Grandbois Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada
Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada

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J Eibl Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada
Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada

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J Crispo Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada
Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada

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D Ansell Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada
Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada

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T C Tai Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada
Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada
Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada
Medical Sciences Division, Departments of Biology, Biomolecular Sciences Program, Northern Ontario School of Medicine, Sudbury, Ontario, Canada

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observed that exposure to high doses of glucocorticoids (GCs) during fetal development, generated either as a consequence of maternal stressors or administered antenatally also reduces birth weight and leads to development of essential hypertension later in

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Thanh Q Dang Department of Biology, Faculty of Science York University, Toronto, Canada

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Nanyoung Yoon Department of Biology, Faculty of Science York University, Toronto, Canada

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Helen Chasiotis Department of Biology, Faculty of Science York University, Toronto, Canada

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Emily C Dunford School of Kinesiology and Health Science, Faculty of Health and Muscle Health Research Center, York University, Toronto, Canada

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Qilong Feng Department of Cellular and Molecular Physiology, College of Medicine, Pennsylvania State University, Hershey, PA, USA

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Pingnian He Department of Cellular and Molecular Physiology, College of Medicine, Pennsylvania State University, Hershey, PA, USA

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Michael C Riddell School of Kinesiology and Health Science, Faculty of Health and Muscle Health Research Center, York University, Toronto, Canada

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Scott P Kelly Department of Biology, Faculty of Science York University, Toronto, Canada

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Gary Sweeney Department of Biology, Faculty of Science York University, Toronto, Canada

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concentrations of the hormone are significantly lower in the target tissue than those in the circulation ( Kolka & Bergman 2012 ). Glucocorticoids are among the most commonly prescribed anti-inflammatory and immunosuppressive medications worldwide ( Clark

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Claire L Wood Division of Developmental Biology, Roslin Institute, University of Edinburgh, Edinburgh, UK

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Ondrej Soucek Department of Paediatrics, 2nd Faculty of Medicine, Charles University in Prague and Motol University Hospital, Prague, Czech Republic
Department of Women’s and Children’s Health, Karolinska Institutet and Pediatric Endocrinology Unit, Karolinska University Hospital, Stockholm, Sweden

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Sze C Wong Developmental Endocrinology Research Group, School of Medicine, University of Glasgow, Glasgow, UK

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Farasat Zaman Department of Women’s and Children’s Health, Karolinska Institutet and Pediatric Endocrinology Unit, Karolinska University Hospital, Stockholm, Sweden

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Colin Farquharson Division of Developmental Biology, Roslin Institute, University of Edinburgh, Edinburgh, UK

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Lars Savendahl Department of Women’s and Children’s Health, Karolinska Institutet and Pediatric Endocrinology Unit, Karolinska University Hospital, Stockholm, Sweden

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S Faisal Ahmed Developmental Endocrinology Research Group, School of Medicine, University of Glasgow, Glasgow, UK

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Introduction and background It is estimated that, at any one time, over 250,000 people are exposed to systemic glucocorticoids (GCs); approximately 10% of children will require GCs at some stage during their childhood ( Mushtaq & Ahmed 2002

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