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Cintia B Ueta Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil
Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Gustavo W Fernandes Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Luciane P Capelo Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Tatiane L Fonseca Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Flávia D'Angelo Maculan Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Cecilia H A Gouveia Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Patrícia C Brum Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Marcelo A Christoffolete Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Marcelo S Aoki Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Carmen L Lancellotti Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Brian Kim Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Antonio C Bianco Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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Miriam O Ribeiro Centro de Ciências Biológicas e da Saúde, Department of Cell and Developmental Biology, School of Arts, Natural and Humans Science Center, Department of Anatomy, Division of Endocrinology, School of Physical Education and Sport, AFIP and Pathology, Universidade Presbiteriana Mackenzie, Rua da Consolação, 869 Prédio 16, 1° Andar, 01302-907 São Paulo, Brazil

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the β 3 receptor ( Granneman 1992 ). Studies performed on isolated brown adipocytes indicate that the β 3 adrenergic receptor plays the leading role in BAT activation ( Zhao et al . 2011 ). However, mice with global β 3 receptor knockout (β 3 KO

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Liping Luo Department of Metabolism and Endocrinology, Metabolic Syndrome Research Center, Key Laboratory of Diabetes Immunology, National Clinical Research Center for Metabolic Diseases, The Second Xiangya Hospital, 
Central South University, Changsha, Hunan, China

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Meilian Liu Department of Metabolism and Endocrinology, Metabolic Syndrome Research Center, Key Laboratory of Diabetes Immunology, National Clinical Research Center for Metabolic Diseases, The Second Xiangya Hospital, 
Central South University, Changsha, Hunan, China
Department of Biochemistry and Molecular Biology, University of New Mexico Health Sciences Center, 
Albuquerque, New Mexico, USA

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elucidate whether transplantation of subcutaneous fat in humans is as efficient as that in rodents. Brown adipocytes were originally thought to be a skeletal muscle-like lineage arising from Myf5 + precursors ( Seale et al . 2008 , Lepper & Fan 2010

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Amy Warner AstraZeneca, Center of Brain, CVMD IMED, Pepparedsleden 1, 43183 Mölndal, Sweden

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Jens Mittag AstraZeneca, Center of Brain, CVMD IMED, Pepparedsleden 1, 43183 Mölndal, Sweden

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contrast to skeletal muscle cells, BAT progenitors initiate the expression of PRDM16 and BMP7 during development, which drives them to become mature brown adipocytes ( Seale et al . 2008 , Tseng et al . 2008 , Rajakumari et al . 2013 ). The most

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Seul Gi Kang Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea
Department of Medical Science, Chungnam National University School of Medicine, Daejeon, Korea

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Hyon-Seung Yi Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea

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Min Jeong Choi Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea
Department of Medical Science, Chungnam National University School of Medicine, Daejeon, Korea

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Min Jeong Ryu Peptron, Inc., Daejeon, Korea

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Saetbyel Jung Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea

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Hyo Kyun Chung Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea

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Joon Young Chang Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea
Department of Medical Science, Chungnam National University School of Medicine, Daejeon, Korea

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Yong Kyung Kim Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea

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Seong Eun Lee Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea
Department of Medical Science, Chungnam National University School of Medicine, Daejeon, Korea

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Hyeon-Woo Kim Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea
Department of Medical Science, Chungnam National University School of Medicine, Daejeon, Korea

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Hoil Choi Department of Biochemistry, Chungnam National University School of Medicine, Daejeon, Korea

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Dong Seok Kim Department of Biochemistry, Chungnam National University School of Medicine, Daejeon, Korea

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Ju Hee Lee Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea

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Koon Soon Kim Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea

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Hyun Jin Kim Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea

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Chul-Ho Lee Animal Model Center, Korea Research Institute of Bioscience and Biotechnology, Daejeon, Korea

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Yuichi Oike Department of Molecular Genetics, Graduate School of Medical Sciences, Kumamoto University, Chuo-ku, Kumamoto, Japan

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Minho Shong Research Center for Endocrine and Metabolic Diseases, Chungnam National University School of Medicine, Daejeon, Korea

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-methylxanthine (0.5 mM), dexamethasone (1 µM), insulin (10 µg/mL) and 10% fetal bovine serum. 3T3-L1 adipocytes, primary hepatocytes, C2C12 myoblasts and mature immortalized brown adipocytes were treated with recombinant angiopoietin-like 6 (ANGPTL6) protein

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Margo P Emont Life Sciences Institute, Department of Molecular and Integrative Physiology, University of Michigan, 210 Washtenaw Avenue, LSI, RM5115A, Ann Arbor, Michigan 48109, USA
Life Sciences Institute, Department of Molecular and Integrative Physiology, University of Michigan, 210 Washtenaw Avenue, LSI, RM5115A, Ann Arbor, Michigan 48109, USA

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Hui Yu Life Sciences Institute, Department of Molecular and Integrative Physiology, University of Michigan, 210 Washtenaw Avenue, LSI, RM5115A, Ann Arbor, Michigan 48109, USA

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Jun Wu Life Sciences Institute, Department of Molecular and Integrative Physiology, University of Michigan, 210 Washtenaw Avenue, LSI, RM5115A, Ann Arbor, Michigan 48109, USA
Life Sciences Institute, Department of Molecular and Integrative Physiology, University of Michigan, 210 Washtenaw Avenue, LSI, RM5115A, Ann Arbor, Michigan 48109, USA

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fat cells have been identified to date ( Rosen & Spiegelman 2014 ). White adipocytes are primarily used for energy storage; they contain a large lipid droplet and few cellular organelles. In contrast, brown adipocytes are primarily a site for adaptive

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Yingning Ji Department of Biochemistry and Molecular Cell Biology, Shanghai Key Laboratory for Tumor Microenvironment and Inflammation, Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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Wei Liu Department of Nuclear Medicine, Fudan University Shanghai Cancer Center, Shanghai, China

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Yemin Zhu Department of Biochemistry and Molecular Cell Biology, Shanghai Key Laboratory for Tumor Microenvironment and Inflammation, Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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Yakui Li Department of Biochemistry and Molecular Cell Biology, Shanghai Key Laboratory for Tumor Microenvironment and Inflammation, Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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Ying Lu Department of Biochemistry and Molecular Biology of School of Basic Medical College of Fudan University, Shanghai, China

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Qi Liu Department of Biochemistry and Molecular Cell Biology, Shanghai Key Laboratory for Tumor Microenvironment and Inflammation, Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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Lingfeng Tong Department of Biochemistry and Molecular Cell Biology, Shanghai Key Laboratory for Tumor Microenvironment and Inflammation, Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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Lei Hu Department of Biochemistry and Molecular Cell Biology, Shanghai Key Laboratory for Tumor Microenvironment and Inflammation, Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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Nannan Xu Department of Biochemistry and Molecular Cell Biology, Shanghai Key Laboratory for Tumor Microenvironment and Inflammation, Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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Zhangbing Chen Department of Biochemistry and Molecular Cell Biology, Shanghai Key Laboratory for Tumor Microenvironment and Inflammation, Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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Na Tian Department of Neurology, Shandong Provincial Hospital Affiliated to Shandong First Medical University, Jinan, Shandong, China

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Lifang Wu Department of Biochemistry and Molecular Cell Biology, Shanghai Key Laboratory for Tumor Microenvironment and Inflammation, Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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Lian Zhu Department of Laboratory Animal Science, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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Shuang Tang Department of Nuclear Medicine, Fudan University Shanghai Cancer Center, Shanghai, China
Cancer Institute, Fudan University Shanghai Cancer Center, Shanghai, China

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Ping Zhang Department of Biochemistry and Molecular Cell Biology, Shanghai Key Laboratory for Tumor Microenvironment and Inflammation, Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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Xuemei Tong Department of Biochemistry and Molecular Cell Biology, Shanghai Key Laboratory for Tumor Microenvironment and Inflammation, Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Shanghai Jiao Tong University School of Medicine, Shanghai, China

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). Cell culture Mouse brown adipocytes (BACs) were provided by Dr Xuyun Zhao in the Department of Biochemistry and Molecular Cell Biology, Shanghai Jiao Tong University School of Medicine ( Wang et al. 2014 ). BACs were cultured in the DMEM medium

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John-Paul Fuller-Jackson
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Belinda A Henry Metabolism, Diabetes and Obesity Program, Monash Biomedicine Discovery Institute, Department of Physiology, Monash University, Clayton, Victoria, Australia

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, mice and rats retain a defined and circumscribed brown fat depot located in the interscapular region ( Sbarbati et al . 1991 , Morroni et al . 1995 ). This contrasts with larger adult mammals, including humans and sheep, where brown adipocytes are

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J Hoppmann
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N Perwitz
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B Meier
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M Fasshauer Department of Internal Medicine I, Department of Internal Medicine III, University of Luebeck, Ratzeburger Allee 160, 23538 Luebeck, Germany

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D Hadaschik
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H Lehnert
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J Klein
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listed as NR3C2 in the MGI Database) is also expressed in white adipocytes as well as in brown adipocytes ( Rondinone et al . 1993 , Zennaro et al . 1998 ). The MR possesses the same affinity for aldosterone and glucocorticoid hormones ( Arriza et al

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Alison Mostyn UP 2012.10.101 EGEAL, School of Veterinary Medicine and Science, INRA UMR 703, INRA, Unité de Recherche 04UR08/03, The Hebrew University of Jerusalem, Unité NOPA, Early Life Research Unit, Institut Polytechnique LaSalle, Beauvais, France

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Linda Attig UP 2012.10.101 EGEAL, School of Veterinary Medicine and Science, INRA UMR 703, INRA, Unité de Recherche 04UR08/03, The Hebrew University of Jerusalem, Unité NOPA, Early Life Research Unit, Institut Polytechnique LaSalle, Beauvais, France

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Thibaut Larcher UP 2012.10.101 EGEAL, School of Veterinary Medicine and Science, INRA UMR 703, INRA, Unité de Recherche 04UR08/03, The Hebrew University of Jerusalem, Unité NOPA, Early Life Research Unit, Institut Polytechnique LaSalle, Beauvais, France

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Samir Dou UP 2012.10.101 EGEAL, School of Veterinary Medicine and Science, INRA UMR 703, INRA, Unité de Recherche 04UR08/03, The Hebrew University of Jerusalem, Unité NOPA, Early Life Research Unit, Institut Polytechnique LaSalle, Beauvais, France

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Pascale Chavatte-Palmer UP 2012.10.101 EGEAL, School of Veterinary Medicine and Science, INRA UMR 703, INRA, Unité de Recherche 04UR08/03, The Hebrew University of Jerusalem, Unité NOPA, Early Life Research Unit, Institut Polytechnique LaSalle, Beauvais, France

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Monia Boukthir UP 2012.10.101 EGEAL, School of Veterinary Medicine and Science, INRA UMR 703, INRA, Unité de Recherche 04UR08/03, The Hebrew University of Jerusalem, Unité NOPA, Early Life Research Unit, Institut Polytechnique LaSalle, Beauvais, France

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Arieh Gertler UP 2012.10.101 EGEAL, School of Veterinary Medicine and Science, INRA UMR 703, INRA, Unité de Recherche 04UR08/03, The Hebrew University of Jerusalem, Unité NOPA, Early Life Research Unit, Institut Polytechnique LaSalle, Beauvais, France

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Jean Djiane UP 2012.10.101 EGEAL, School of Veterinary Medicine and Science, INRA UMR 703, INRA, Unité de Recherche 04UR08/03, The Hebrew University of Jerusalem, Unité NOPA, Early Life Research Unit, Institut Polytechnique LaSalle, Beauvais, France

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Michael E Symonds UP 2012.10.101 EGEAL, School of Veterinary Medicine and Science, INRA UMR 703, INRA, Unité de Recherche 04UR08/03, The Hebrew University of Jerusalem, Unité NOPA, Early Life Research Unit, Institut Polytechnique LaSalle, Beauvais, France

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Latifa Abdennebi-Najar UP 2012.10.101 EGEAL, School of Veterinary Medicine and Science, INRA UMR 703, INRA, Unité de Recherche 04UR08/03, The Hebrew University of Jerusalem, Unité NOPA, Early Life Research Unit, Institut Polytechnique LaSalle, Beauvais, France

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brown adipocytes in perirenal adipose tissue ( Figs 2 and 3 ), whereas UCP1 was not present in the subcutaneous depot. These brown adipocytes showed two distinct morphologies, being either large and shapeless or with a more typical multilocular

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Yang Chen Transgenic Research Center, Northeast Normal University, Changchun, China

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Mingyue Zhao Transgenic Research Center, Northeast Normal University, Changchun, China

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Chenhao Wang Transgenic Research Center, Northeast Normal University, Changchun, China

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Huaizhen Wen Transgenic Research Center, Northeast Normal University, Changchun, China

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Yuntao Zhang Transgenic Research Center, Northeast Normal University, Changchun, China

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Mingxu Lu Transgenic Research Center, Northeast Normal University, Changchun, China

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Salah Adlat Transgenic Research Center, Northeast Normal University, Changchun, China

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Tingting Zheng Transgenic Research Center, Northeast Normal University, Changchun, China

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Mingjiao Zhang Transgenic Research Center, Northeast Normal University, Changchun, China

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Dan Li Transgenic Research Center, Northeast Normal University, Changchun, China

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Xiaodan Lu Transgenic Research Center, Northeast Normal University, Changchun, China

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Mengwei Guo Transgenic Research Center, Northeast Normal University, Changchun, China

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Hongyu Chen Transgenic Research Center, Northeast Normal University, Changchun, China

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Luqing Zhang Transgenic Research Center, Northeast Normal University, Changchun, China
Key Laboratory of Molecular Epigenetics of Ministry of Education, Northeast Normal University, Changchun, China

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Xuechao Feng Transgenic Research Center, Northeast Normal University, Changchun, China
Key Laboratory of Molecular Epigenetics of Ministry of Education, Northeast Normal University, Changchun, China

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Yaowu Zheng Transgenic Research Center, Northeast Normal University, Changchun, China
Key Laboratory of Molecular Epigenetics of Ministry of Education, Northeast Normal University, Changchun, China

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, Kershaw & Flier 2004 ). Brown adipocytes, in contrast, originate from precursors that express Myf5 ( Kopelman 2000 ) and contain a huge number of mitochondria and multiple small lipid droplets for efficient energy use. BAT is rich in vasculature and

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