The role of skeletal muscle Akt in the regulation of muscle mass and glucose homeostasis

DeFronzo, 1981, The effect of insulin on the disposal of intravenous glucose. Results from indirect calorimetry and hepatic and femoral venous catheterization∖Effects of insulin on peripheral and splanchnic glucose metabolism in noninsulin-dependent (type II) diabetes m, Diabetes, 30, 1000, 10.2337/diab.30.12.1000

DeFronzo, 2004, Pathogenesis of type 2 diabetes mellitus, Medical Clinics of North America, 10.1016/j.mcna.2004.04.013

Kelley, 1994, Impaired postprandial glucose utilization in non-insulin-dependent diabetes mellitus, Metabolism, 43, 1549, 10.1016/0026-0495(94)90015-9

Kelley, 1988, Skeletal muscle glycolysis, oxidation, and storage of an oral glucose load, Journal of Clinical Investigation, 10.1172/JCI113489

Taylor, 1996, Direct assessment of liver glycogen storage by 13C nuclear magnetic resonance spectroscopy and regulation of glucose homeostasis after a mixed meal in normal subjects, Journal of Clinical Investigation, 10.1172/JCI118379

DeFronzo, 2009, Skeletal muscle insulin resistance is the primary defect in type 2 diabetes, Diabetes Care, 32, S157, 10.2337/dc09-S302

Whiteman, 2002, Role of Akt/protein kinase B in metabolism, Trends in Endocrinology and Metabolism TEM, 13, 444, 10.1016/S1043-2760(02)00662-8

Peng, 2003, Dwarfism, impaired skin development, skeletal muscle atrophy, delayed bone development, and impeded adipogenesis in mice lacking Akt1 and Akt2, Genes and Development, 17, 1352, 10.1101/gad.1089403

Easton, 2005, Role for Akt3/protein kinase B in attainment of normal brain size, Molecular and Cellular Biology, 25, 1869, 10.1128/MCB.25.5.1869-1878.2005

Yang, 2003, Protein kinase Bα/Akt1 regulates placental development and fetal growth, Journal of Biological Chemistry, 278, 32124, 10.1074/jbc.M302847200

Cho, 2001, Akt1/PKBα is required for normal growth but dispensable for maintenance of glucose homeostasis in mice, Journal of Biological Chemistry, 276, 38349, 10.1074/jbc.C100462200

Cho, 2001, Insulin resistance and a diabetes mellitus-like syndrome in mice lacking the protein kinase Akt2 (PKBbeta), Science, 292, 1728, 10.1126/science.292.5522.1728

Garofalo, 2003, Severe diabetes, age-dependent loss of adipose tissue, and mild growth deficiency in mice lacking Akt2/PKBβ, Journal of Clinical Investigation, 112, 197, 10.1172/JCI16885

Lu, 2012, Insulin regulates liver metabolism in vivo in the absence of hepatic Akt and Foxo1, Nature Medicine, 18, 388, 10.1038/nm.2686

Krook, 1998, Insulin-stimulated Akt kinase activity is reduced in skeletal muscle from NIDDM subjects, Diabetes, 47, 1281, 10.2337/diab.47.8.1281

Zierath, 1998, Insulin action in skeletal muscle from patients with NIDDM, Molecular and Cellular Biochemistry, 182, 153, 10.1023/A:1006861628496

Højlund, 2009, Dysregulation of glycogen synthase COOH- and NH2-terminal phosphorylation by insulin in obesity and type 2 diabetes mellitus, Journal of Clinical Endocrinology and Metabolism, 94, 4547, 10.1210/jc.2009-0897

Brozinick, 2003, Defective signaling through Akt-2 and -3 but not Akt-1 in insulin-resistant human skeletal muscle: potential role in insulin resistance, Diabetes, 52, 935, 10.2337/diabetes.52.4.935

Cozzone, 2008, Isoform-specific defects of insulin stimulation of Akt/protein kinase B (PKB) in skeletal muscle cells from type 2 diabetic patients, Diabetologia, 51, 512, 10.1007/s00125-007-0913-8

Karlsson, 2005, Insulin-stimulated phosphorylation of the Akt substrate AS160 is impaired in skeletal muscle of type 2 diabetic subjects, Diabetes, 54, 1692, 10.2337/diabetes.54.6.1692

Tremblay, 2001, Defective insulin-induced GLUT4 translocation in skeletal muscle of high fat-fed rats is associated with alterations in both Akt/protein kinase B and atypical protein kinase C (ζ/λ) activities, Diabetes, 50, 1901, 10.2337/diabetes.50.8.1901

Shao, 2000, Decreased Akt kinase activity and insulin resistance C57BL/KsJ-Lepr(db/db) mice, Journal of Endocrinology, 167, 107, 10.1677/joe.0.1670107

Lai, 2012, A novel PKB/Akt inhibitor, MK-2206, effectively inhibits insulin-stimulated glucose metabolism and protein synthesis in isolated rat skeletal muscle, Biochemical Journal, 447, 137, 10.1042/BJ20120772

Yeh, 1995, The effects of wortmannin on rat skeletal muscle. Dissociation of signaling pathways for insulin- and contraction-activated hexose transport, Journal of Biological Chemistry, 270, 2107, 10.1074/jbc.270.5.2107

Ducommun, 2012, Thr649Ala-AS160 knock-in mutation does not impair contraction/AICAR-induced glucose transport in mouse muscle, American Journal of Physiology Endocrinology and Metabolism, 302, E1036, 10.1152/ajpendo.00379.2011

Wang, 2013, AS160 deficiency causes whole-body insulin resistance via composite effects in multiple tissues, Biochemical Journal, 449, 479, 10.1042/BJ20120702

Chen, 2011, Mice with as160/TBC1D4-Thr649Ala Knockin mutation are glucose intolerant with reduced insulin sensitivity and altered GLUT4 trafficking, Cell Metabolism, 13, 68, 10.1016/j.cmet.2010.12.005

Sakamoto, 2008, Emerging role for AS160/TBC1D4 and TBC1D1 in the regulation of GLUT4 traffic, American Journal of Physiology-Endocrinology and Metabolism, 295, E29, 10.1152/ajpendo.90331.2008

Whitfield, 2017, Ablating the protein TBC1D1 impairs contraction-induced sarcolemmal glucose transporter 4 redistribution but not insulin-mediated responses in rats, Journal of Biological Chemistry, 292, 16653, 10.1074/jbc.M117.806786

O'Neill, 2015, Differential role of insulin/IGF-1 receptor signaling in muscle growth and glucose homeostasis, Cell Reports, 11, 1220, 10.1016/j.celrep.2015.04.037

Geddis, 2017, AKT2 is the predominant Akt isoform expressed in human skeletal muscle, The FASEB Journal, 31

Wan, 2012, Loss of Akt1 in mice increases energy expenditure and protects against diet-induced obesity, Molecular and Cellular Biology, 32, 96, 10.1128/MCB.05806-11

Green, 2008, Use of Akt inhibitor and a drug-resistant mutant validates a critical role for protein kinase B/Akt in the insulin-dependent regulation of glucose and system A amino acid uptake, Journal of Biological Chemistry, 283, 27653, 10.1074/jbc.M802623200

McCarthy, 2012, Inducible Cre transgenic mouse strain for skeletal muscle-specific gene targeting, Skeletal Muscle, 2, 8, 10.1186/2044-5040-2-8

Kjøbsted, 2018, AMPK in skeletal muscle function and metabolism, FASEB Journal, 10.1096/fj.201700442R

Fujii, 2004, Regulation of glucose transport by the AMP-activated protein kinase, Proceedings of the Nutrition Society, 63, 205, 10.1079/PNS2004340

Funai, 2009, Inhibition of contraction-stimulated amp-activated protein kinase inhibits contraction-stimulated increases in pas-tbc1d1 and glucose transport without altering pas-as160 in rat skeletal muscle, Diabetes, 58, 1096, 10.2337/db08-1477

Mu, 2001, A role for AMP-activated protein kinase in contraction- and hypoxia-regulated glucose transport in skeletal muscle, Molecular Cell, 7, 1085, 10.1016/S1097-2765(01)00251-9

Cokorinos, 2017, Activation of skeletal muscle AMPK promotes glucose disposal and glucose lowering in non-human primates and mice, Cell Metabolism, 25, 1147, 10.1016/j.cmet.2017.04.010

Steinberg, 2006, Tumor necrosis factor α-induced skeletal muscle insulin resistance involves suppression of AMP-kinase signaling, Cell Metabolism, 4, 465, 10.1016/j.cmet.2006.11.005

Bandyopadhyay, 2006, Increased malonyl-CoA levels in muscle from obese and type 2 diabetic subjects lead to decreased fatty acid oxidation and increased lipogenesis; thiazolidinedione treatment reverses these defects, Diabetes, 55, 2277, 10.2337/db06-0062

Geng, 2018, H19 lncRNA promotes skeletal muscle insulin sensitivity in part by targeting AMPK, Diabetes, 67, 2183, 10.2337/db18-0370

Sriwijitkamol, 2007, Effect of acute exercise on AMPK signaling in skeletal muscle of subjects with type 2 diabetes: a time-course and dose-response study, Diabetes, 10.2337/db06-1119

Brozinick, 1998, Insulin, but not contraction, activates Akt/PKB in isolated rat skeletal muscle, Journal of Biological Chemistry, 273, 14679, 10.1074/jbc.273.24.14679

Kjøbsted, 2017, Enhanced muscle insulin sensitivity after contraction/exercise is mediated by AMPK, Diabetes, 10.2337/db16-0530

Hamada, 2006, Increased submaximal insulin-stimulated glucose uptake in mouse skeletal muscle after treadmill exercise, Journal of Applied Physiology, 10.1152/japplphysiol.00416.2006

Holloszy, 2005, Exercise-induced increase in muscle insulin sensitivity, Journal of Applied Physiology, 10.1152/japplphysiol.00123.2005

Ching, 2010, A role for AMPK in increased insulin action after serum starvation, American Journal of Physiology Cell Physiology, 10.1152/ajpcell.00514.2009

Smith, 2005, AICAR and hyperosmotic stress increase insulin-stimulated glucose transport, Journal of Applied Physiology, 10.1152/japplphysiol.01297.2004

Fisher, 2017, Activation of AMP kinase enhances sensitivity of muscle glucose transport to insulin, American Journal of Physiology Endocrinology and Metabolism

Bain, 2007, The selectivity of protein kinase inhibitors: a further update, Biochemical Journal, 10.1042/BJ20070797

Dite, 2018, AMP-activated protein kinase selectively inhibited by the type II inhibitor SBI-0206965, Journal of Biological Chemistry, 10.1074/jbc.RA118.003547

Samuel, 2016, The pathogenesis of insulin resistance: integrating signaling pathways and substrate flux, Journal of Clinical Investigation, 126, 12, 10.1172/JCI77812

Long, 2011, Insulin receptor substrates Irs1 and Irs2 coordinate skeletal muscle growth and metabolism via the Akt and AMPK pathways, Molecular and Cellular Biology, 31, 430, 10.1128/MCB.00983-10

Luo, 2006, Loss of class IA PI3K signaling in muscle leads to impaired muscle growth, insulin response, and hyperlipidemia, Cell Metabolism, 3, 355, 10.1016/j.cmet.2006.04.003

O'Neill, 2016, Insulin and IGF-1 receptors regulate FoxO-mediated signaling in muscle proteostasis, Journal of Clinical Investigation, 126, 3433, 10.1172/JCI86522

Kjøbsted, 2015, Prior AICAR stimulation increases insulin sensitivity in mouse skeletal muscle in an AMPK-dependent manner, Diabetes, 10.2337/db14-1402

Steinberg, 2006, Reduced glycogen availability is associated with increased AMPKα2 activity, nuclear AMPKα2 protein abundance, and GLUT4 mRNA expression in contracting human skeletal muscle, Applied Physiology Nutrition and Metabolism, 10.1139/h06-003

McBride, 2009, The glycogen-binding domain on the AMPK β subunit Allows the kinase to act as a glycogen sensor, Cell Metabolism, 10.1016/j.cmet.2008.11.008

Wojtaszewski, 2002, Dissociation of AMPK activity and ACCβ phosphorylation in human muscle during prolonged exercise, Biochemical and Biophysical Research Communications, 10.1016/S0006-291X(02)02465-8

Taylor, 2008, Discovery of TBC1D1 as an insulin-, AICAR-, and contraction-stimulated signaling nexus in mouse skeletal muscle, Journal of Biological Chemistry, 283, 9787, 10.1074/jbc.M708839200

Rudich, 2013, Putting Rac1 on the path to glucose uptake, Diabetes, 10.2337/db13-0381

Sylow, 2014, Akt and Rac1 signaling are jointly required for insulin-stimulated glucose uptake in skeletal muscle and downregulated in insulin resistance, Cellular Signalling, 26, 323, 10.1016/j.cellsig.2013.11.007

Ueda, 2010, Crucial role of the small GTPase Rac1 in insulin-stimulated translocation of glucose transporter 4 to the mouse skeletal muscle sarcolemma, The FASEB Journal, 10.1096/fj.09-137380

Sylow, 2013, Rac1 signaling is required for insulin-stimulated glucose uptake and is dysregulated in insulin-resistant murine and human skeletal muscle, Diabetes, 62, 1865, 10.2337/db12-1148

Tunduguru, 2014, Signaling of the p21-activated kinase (PAK1) coordinates insulin-stimulated actin remodeling and glucose uptake in skeletal muscle cells, Biochemical Pharmacology, 10.1016/j.bcp.2014.08.033

Moller, 2019, The role of p-21 activated kinases (PAKs) in glucose homeostasis and skeletal muscle glucose uptake, BioRxiv

Nozaki, 2013, Akt2 regulates Rac1 activity in the insulin-dependent signaling pathway leading to GLUT4 translocation to the plasma membrane in skeletal muscle cells, Cellular Signalling, 10.1016/j.cellsig.2013.02.023

Takenaka, 2014, Role of the guanine nucleotide exchange factor in Akt2-mediated plasma membrane translocation of GLUT4 in insulin-stimulated skeletal muscle, Cellular Signalling, 10.1016/j.cellsig.2014.07.002

Takenaka, 2019, Involvement of the protein kinase Akt2 in insulin-stimulated Rac1 activation leading to glucose uptake in mouse skeletal muscle, PLoS One, 10.1371/journal.pone.0212219

Titchenell, 2016, Direct hepatocyte insulin signaling is required for lipogenesis but is dispensable for the suppression of glucose production, Cell Metabolism, 23, 1154, 10.1016/j.cmet.2016.04.022

McKeel, 1970, Preparation and characterization of a plasma membrane fraction from isolated fat cells, Journal of Cell Biology, 10.1083/jcb.44.2.417

Tonks, 2013, Impaired Akt phosphorylation in insulin-resistant human muscle is accompanied by selective and heterogeneous downstream defects, Diabetologia, 56, 875, 10.1007/s00125-012-2811-y

Fazakerley, 2018, Muscle and adipose tissue insulin resistance: malady without mechanism?, Journal of Lipid Research, 10.1194/jlr.R087510

Hoehn, 2008, IRS1-independent defects define major nodes of insulin resistance, Cellular Metabolism, 7, 421, 10.1016/j.cmet.2008.04.005