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Caishun Zhang Special Medicine Department, College of Basic Medicine, Qingdao University, Qingdao, China

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Junhua Yuan Special Medicine Department, College of Basic Medicine, Qingdao University, Qingdao, China

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Qian Lin Special Medicine Department, College of Basic Medicine, Qingdao University, Qingdao, China

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Manwen Li Special Medicine Department, College of Basic Medicine, Qingdao University, Qingdao, China

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Liuxin Wang Hyperbaric Oxygen Therapy Department, Yuhuangding Hospital Affiliated to Qingdao University, Yantai, China

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Rui Wang Special Medicine Department, College of Basic Medicine, Qingdao University, Qingdao, China

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Xi Chen Physiology Department, College of Basic Medicine, Qingdao University, Qingdao, China

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Zhengyao Jiang Physiology Department, College of Basic Medicine, Qingdao University, Qingdao, China

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Kun Zhu Intensive Care Unit Department, Affiliated Hospital of Qingdao University, Qingdao, China

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Xiaoli Chang Institute of Acupuncture, Shandong University of Traditional Chinese Medicine, Jinan, China

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Bin Wang Special Medicine Department, College of Basic Medicine, Qingdao University, Qingdao, China
Medical Microbiology Department, College of Basic Medicine, Qingdao University, Qingdao, China

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Jing Dong Special Medicine Department, College of Basic Medicine, Qingdao University, Qingdao, China
Physiology Department, College of Basic Medicine, Qingdao University, Qingdao, China

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stimulate the release of growth hormone from the pituitary ( 1 , 2 , 3 ). Then it was reported to be also involved in the regulation of feeding ( 4 ), body weight ( 5 ), and energy metabolism ( 6 ). A key role had been established for ghrelin in energy

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Michaela Keuper Department of Molecular Biosciences, The Wenner-Gren Institute, Stockholm University, Stockholm, Sweden

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aspects of MΦ-induced browning have been questioned ( 29 , 30 ), but most studies collectively support a role for MΦ in the energy metabolism of adipocytes, in particular controlling adipocyte mitochondrial function ( 26 , 27 , 31 , 32 , 33 , 34

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Carlijn A Hoekx Division of Endocrinology, Department of Medicine, Leiden University Medical Center, Leiden, The Netherlands
Einthoven Laboratory for Experimental Vascular Medicine, Leiden University Medical Center, Leiden, The Netherlands

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Borja Martinez-Tellez Division of Endocrinology, Department of Medicine, Leiden University Medical Center, Leiden, The Netherlands
Einthoven Laboratory for Experimental Vascular Medicine, Leiden University Medical Center, Leiden, The Netherlands
Department of Nursing Physiotherapy and Medicine, SPORT Research Group (CTS-1024), CERNEP Research Center, University of Almería, Almería, Spain
Biomedical Research Unit, Torrecárdenas University Hospital, Almería, Spain
CIBER de Fisiopatología de la Obesidad y Nutrición (CIBEROBN), Instituto de Salud Carlos III, Granada, Spain

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Maaike E Straat Division of Endocrinology, Department of Medicine, Leiden University Medical Center, Leiden, The Netherlands
Einthoven Laboratory for Experimental Vascular Medicine, Leiden University Medical Center, Leiden, The Netherlands

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Magdalena M A Verkleij Division of Endocrinology, Department of Medicine, Leiden University Medical Center, Leiden, The Netherlands
Einthoven Laboratory for Experimental Vascular Medicine, Leiden University Medical Center, Leiden, The Netherlands

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Mirjam Kemmeren Division of Endocrinology, Department of Medicine, Leiden University Medical Center, Leiden, The Netherlands
Einthoven Laboratory for Experimental Vascular Medicine, Leiden University Medical Center, Leiden, The Netherlands

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Sander Kooijman Division of Endocrinology, Department of Medicine, Leiden University Medical Center, Leiden, The Netherlands
Einthoven Laboratory for Experimental Vascular Medicine, Leiden University Medical Center, Leiden, The Netherlands

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Martin Uhrbom Bioscience Metabolism, Research and Early Development, Cardiovascular, Renal and Metabolism (CVRM), BioPharmaceuticals R&D, AstraZeneca Gothenburg, Sweden
Department of Medicine, Huddinge, Karolinska Institutet Campus Flemingsberg, Neo Building, Huddinge, Sweden

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Saskia C A de Jager Laboratory of Experimental Cardiology, University Medical Center Utrecht, Utrecht, The Netherlands

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Patrick C N Rensen Division of Endocrinology, Department of Medicine, Leiden University Medical Center, Leiden, The Netherlands
Einthoven Laboratory for Experimental Vascular Medicine, Leiden University Medical Center, Leiden, The Netherlands

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Mariëtte R Boon Division of Endocrinology, Department of Medicine, Leiden University Medical Center, Leiden, The Netherlands
Einthoven Laboratory for Experimental Vascular Medicine, Leiden University Medical Center, Leiden, The Netherlands

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that could enhance energy metabolism ( 15 ). Preclinical experiments have identified two batokines that have a prominent effect on energy balance: fibroblast growth factor 21 (FGF21) and growth differentiation factor 15 (GDF15) ( 16 , 17 ). FGF21, when

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Selina Mäkinen Minerva Foundation Institute for Medical Research, Helsinki, Finland
Department of Medicine, University of Helsinki, Helsinki University Central Hospital, Helsinki, Finland

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Neeta Datta Minerva Foundation Institute for Medical Research, Helsinki, Finland
Department of Medicine, University of Helsinki, Helsinki University Central Hospital, Helsinki, Finland

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Yen H Nguyen Minerva Foundation Institute for Medical Research, Helsinki, Finland
Department of Medicine, University of Helsinki, Helsinki University Central Hospital, Helsinki, Finland

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Petro Kyrylenko Minerva Foundation Institute for Medical Research, Helsinki, Finland
Department of Medicine, University of Helsinki, Helsinki University Central Hospital, Helsinki, Finland

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Markku Laakso Institute of Clinical Medicine, Internal Medicine, University of Eastern Finland, Kuopio, Finland

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Heikki A Koistinen Minerva Foundation Institute for Medical Research, Helsinki, Finland
Department of Medicine, University of Helsinki, Helsinki University Central Hospital, Helsinki, Finland

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Objectives

Simvastatin use is associated with muscular side effects, and increased risk for type 2 diabetes (T2D). In clinical use, simvastatin is administered in inactive lipophilic lactone-form, which is then converted to active acid-form in the body. Here, we have investigated if lactone- and acid-form simvastatin differentially affect glucose metabolism and mitochondrial respiration in primary human skeletal muscle cells.

Methods

Muscle cells were exposed separately to lactone- and acid-form simvastatin for 48 h. After pre-exposure, glucose uptake and glycogen synthesis were measured using radioactive tracers; insulin signalling was detected with Western blotting; and glycolysis, mitochondrial oxygen consumption and ATP production were measured with Seahorse XFe96 analyzer.

Results

Lactone-form simvastatin increased glucose uptake and glycogen synthesis, whereas acid-form simvastatin did not affect glucose uptake and decreased glycogen synthesis. Phosphorylation of insulin signalling targets Akt substrate 160 kDa (AS160) and glycogen synthase kinase 3β (GSK3β) was upregulated with lactone-, but not with acid-form simvastatin. Exposure to both forms of simvastatin led to a decrease in glycolysis and glycolytic capacity, as well as to a decrease in mitochondrial respiration and ATP production.

Conclusions

These data suggest that lactone- and acid-forms of simvastatin exhibit differential effects on non-oxidative glucose metabolism as lactone-form increases and acid-form impairs glucose storage into glycogen, suggesting impaired insulin sensitivity in response to acid-form simvastatin. Both forms profoundly impair oxidative glucose metabolism and energy production in human skeletal muscle cells. These effects may contribute to muscular side effects and risk for T2D observed with simvastatin use.

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Selina Mäkinen Minerva Foundation Institute for Medical Research, Helsinki, Finland
Department of Medicine, University of Helsinki, Helsinki University Central Hospital, Helsinki, Finland

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Neeta Datta Minerva Foundation Institute for Medical Research, Helsinki, Finland
Department of Medicine, University of Helsinki, Helsinki University Central Hospital, Helsinki, Finland

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Yen H Nguyen Minerva Foundation Institute for Medical Research, Helsinki, Finland
Department of Medicine, University of Helsinki, Helsinki University Central Hospital, Helsinki, Finland

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Petro Kyrylenko Minerva Foundation Institute for Medical Research, Helsinki, Finland
Department of Medicine, University of Helsinki, Helsinki University Central Hospital, Helsinki, Finland

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Markku Laakso Institute of Clinical Medicine, Internal Medicine, University of Eastern Finland, Kuopio, Finland

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Heikki A Koistinen Minerva Foundation Institute for Medical Research, Helsinki, Finland
Department of Medicine, University of Helsinki, Helsinki University Central Hospital, Helsinki, Finland

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Jia Liu Department of Endocrinology, Beijing Chao-yang Hospital, Capital Medical University, Chaoyang District, Beijing, China

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Lin Zhang Department of Endocrinology, Beijing Chao-yang Hospital, Capital Medical University, Chaoyang District, Beijing, China

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Jing Fu Department of Endocrinology, Beijing Chao-yang Hospital, Capital Medical University, Chaoyang District, Beijing, China

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Qiu Wang Department of Endocrinology, Beijing Chao-yang Hospital, Capital Medical University, Chaoyang District, Beijing, China

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Guang Wang Department of Endocrinology, Beijing Chao-yang Hospital, Capital Medical University, Chaoyang District, Beijing, China

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from anterior pituitary and named for its crucial effect on lactation ( 8 , 9 ). Recent studies have demonstrated that PRL plays an important role in regulating energy metabolism ( 8 , 9 , 10 , 11 , 12 , 13 , 14 , 15 , 16 ). PRL administration

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Florian W Kiefer Clinical Division of Endocrinology and Metabolism, Department of Medicine III, Medical University of Vienna, Vienna, Austria

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that results in increased energy wasting and body weight loss in humans is yet to be shown. Function of human BAT: relevance for energy metabolism Soon after the detection of cold-induced BAT in human adults in 2009, numerous follow-up studies

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Jeyanthini Risikesan Department of Endocrinology and Internal Medicine, Aarhus University Hospital, Aarhus, Denmark

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Birgitte Nellemann Department of Endocrinology and Internal Medicine, Aarhus University Hospital, Aarhus, Denmark
Department of Rheumatology, Diakonhjemmet Hospital, Oslo, Norway

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Britt Christensen Department of Endocrinology and Internal Medicine, Aarhus University Hospital, Aarhus, Denmark

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Jens Otto Lunde Jørgensen Department of Endocrinology and Internal Medicine, Aarhus University Hospital, Aarhus, Denmark

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Søren Nielsen Department of Endocrinology and Internal Medicine, Aarhus University Hospital, Aarhus, Denmark
Steno Diabetes Center Aarhus, Aarhus University Hospital, Aarhus, Denmark

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, 9 ). Other studies indicate that EPO has effects on lipid and energy metabolism. However, the mechanisms behind these effects remain unclear ( 10 , 11 , 12 , 13 ). Treatment with rHuEpo has been reported to improve insulin sensitivity and

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Tao Mei Department of Neurosurgery, Fuzhou General Hospital, Fuzhou, China
Fuzong Clinical Medical College of Fujian Medical University, Fuzhou, China

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Jianhe Zhang Department of Neurosurgery, Fuzhou General Hospital, Fuzhou, China

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Liangfeng Wei Department of Neurosurgery, Fuzhou General Hospital, Fuzhou, China

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Xingfeng Qi Department of Pathology, Fuzhou General Hospital, Fuzhou, China

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Yiming Ma Department of Neurosurgery, Liuzhou General Hospital, Liuzhou, China

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Xianhua Liu Department of Pathology, Fuzhou General Hospital, Fuzhou, China

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Shaohua Chen Department of Pathology, The Second Affiliated Hospital of Fujian Medical University, Quanzhou, China

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Songyuan Li Department of Neurosurgery, Fuzhou General Hospital, Fuzhou, China

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Jianwu Wu Department of Neurosurgery, Fuzhou General Hospital, Fuzhou, China

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Shousen Wang Department of Neurosurgery, Fuzhou General Hospital, Fuzhou, China

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hypoxia ( 3 ). Because glucose is the main source of energy for tumor cells, study of the glucose metabolism in PA will improve our understanding of the biobehavioral mechanisms of PA cell energy metabolism. While glucose is the main energy source of the

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L Johnsen Department of Large Animal Sciences, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

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N B Lyckegaard Department of Large Animal Sciences, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

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P Khanal Department of Large Animal Sciences, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

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B Quistorff Department of Biomedical Sciences, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

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K Raun Diabetes and Obesity Pharmacology, Novo Nordisk A/S, Måløv, Denmark

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M O Nielsen Department of Large Animal Sciences, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

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downstream regulators ( 45 , 24 ). This interference of peripheral TH function in tissues, quantitatively important in energy metabolism, is in line with the observation that HIGH:CONV sheep did not increase their heart rate in response to a thyroxine surge

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