{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,17]],"date-time":"2026-10-17T00:45:52Z","timestamp":1792197952532,"version":"4.4.0"},"reference-count":177,"publisher":"MDPI AG","issue":"15","license":[{"start":{"date-parts":[[2021,7,30]],"date-time":"2021-07-30T00:00:00Z","timestamp":1627603200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["RGPIN-2021-03724"],"award-info":[{"award-number":["RGPIN-2021-03724"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Canadian Institutes of Health Research","award":["MOP-93760"],"award-info":[{"award-number":["MOP-93760"]}]},{"name":"Canadian Institutes of Health Research","award":["MOV-409262"],"award-info":[{"award-number":["MOV-409262"]}]},{"name":"Spanish Ministry of Science and Innovation","award":["FIS 18\/00075"],"award-info":[{"award-number":["FIS 18\/00075"]}]},{"name":"Spanish Ministry of Science and Innovation","award":["FIS 21\/00215"],"award-info":[{"award-number":["FIS 21\/00215"]}]},{"name":"Sociedad Espa\u00f1ola de Neumolog\u00eda y Cirug\u00eda Tor\u00e1cica (SEPAR)","award":["SEPAR 2020"],"award-info":[{"award-number":["SEPAR 2020"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>The maintenance of mitochondrial integrity is critical for muscle health. Mitochondria, indeed, play vital roles in a wide range of cellular processes, including energy supply, Ca2+ homeostasis, retrograde signaling, cell death, and many others. All mitochondria-containing cells, including skeletal muscle cells, dispose of several pathways to maintain mitochondrial health, including mitochondrial biogenesis, mitochondrial-derived vesicles, mitochondrial dynamics (fusion and fission process shaping mitochondrial morphology), and mitophagy\u2014the process in charge of the removal of mitochondria though autophagy. The loss of skeletal muscle mass (atrophy) is a major health problem worldwide, especially in older people. Currently, there is no treatment to counteract the progressive decline in skeletal muscle mass and strength that occurs with aging, a process termed sarcopenia. There is increasing data, including our own, suggesting that accumulation of dysfunctional mitochondria contributes to the development of sarcopenia. Impairments in mitochondrial dynamics and mitophagy were recently proposed to contribute to sarcopenia. This review summarizes the current state of knowledge on the role played by mitochondrial dynamics and mitophagy in skeletal muscle health and in the development of sarcopenia. We also highlight recent studies showing that enhancing mitophagy in skeletal muscle is a promising therapeutic target to prevent or even treat skeletal muscle dysfunction in the elderly.<\/jats:p>","DOI":"10.3390\/ijms22158179","type":"journal-article","created":{"date-parts":[[2021,7,30]],"date-time":"2021-07-30T03:46:42Z","timestamp":1627616802000},"page":"8179","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":274,"title":["Mitochondrial Dynamics and Mitophagy in Skeletal Muscle Health and Aging"],"prefix":"10.3390","volume":"22","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7637-5445","authenticated-orcid":false,"given":"Jean-Philippe","family":"Leduc-Gaudet","sequence":"first","affiliation":[{"name":"Research Institute of the McGill University Health Centre, Department of Critical Care, Montr\u00e9al, QC H4A 3J1, Canada"},{"name":"Meakins-Christie Laboratories, Department of Medicine, McGill University, Montr\u00e9al, QC H4A 3J1, Canada"},{"name":"D\u00e9partement des Sciences de l\u2019activit\u00e9 Physique, Facult\u00e9 des Sciences, UQAM, Montr\u00e9al, QC H2X 1Y4, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sabah N. A.","family":"Hussain","sequence":"additional","affiliation":[{"name":"Research Institute of the McGill University Health Centre, Department of Critical Care, Montr\u00e9al, QC H4A 3J1, Canada"},{"name":"Meakins-Christie Laboratories, Department of Medicine, McGill University, Montr\u00e9al, QC H4A 3J1, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Esther","family":"Barreiro","sequence":"additional","affiliation":[{"name":"Pulmonology Department-Muscle Wasting & Cachexia in Chronic Respiratory Diseases & Lung Cancer Research Group, IMIM-Hospital del Mar, Parc de Salut Mar, Biomedical Research Park (PRBB), C\/Dr. Aiguader, 88, 08003 Barcelona, Spain"},{"name":"Centro de Investigaci\u00f3n en Red de Enfermedades Respiratorias (CIBERES), Instituto de Salud Carlos III (ISCIII), 28029 Madrid, Spain"},{"name":"Health and Experimental Sciences Department (CEXS), Pompeu Fabra University (UPF), Biomedical Research Park (PRBB), C\/Dr. Aiguader, 88, 08003 Barcelona, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8543-3619","authenticated-orcid":false,"given":"Gilles","family":"Gouspillou","sequence":"additional","affiliation":[{"name":"Research Institute of the McGill University Health Centre, Department of Critical Care, Montr\u00e9al, QC H4A 3J1, Canada"},{"name":"Meakins-Christie Laboratories, Department of Medicine, McGill University, Montr\u00e9al, QC H4A 3J1, Canada"},{"name":"D\u00e9partement des Sciences de l\u2019activit\u00e9 Physique, Facult\u00e9 des Sciences, UQAM, Montr\u00e9al, QC H2X 1Y4, Canada"},{"name":"Centre de Recherche de l\u2019Institut Universitaire de G\u00e9riatrie de Montr\u00e9al, Montr\u00e9al, QC H3W 1W5, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,7,30]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"330","DOI":"10.1038\/s41467-020-20123-1","article-title":"Mechanisms of muscle atrophy and hypertrophy: Implications in health and disease","volume":"12","author":"Sartori","year":"2021","journal-title":"Nat. Commun."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1007\/s12603-020-1325-4","article-title":"Sarcopenia in Community-Dwelling Oldest Old Is Associated with Disability and Poor Physical Function","volume":"24","author":"Xu","year":"2020","journal-title":"J. Nutr. Health Aging"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1002\/jcsm.12160","article-title":"Sarcopenia and physical independence in older adults: The independent and synergic role of muscle mass and muscle function","volume":"8","author":"Cyrino","year":"2017","journal-title":"J. Cachexia Sarcopenia Muscle"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1016\/j.asjsur.2019.10.010","article-title":"Impact of sarcopenia in trauma and surgical patient population: A literature review","volume":"43","author":"Wahlen","year":"2020","journal-title":"Asian J. Surg."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Zhang, X., Zhang, W., Wang, C., Tao, W., Dou, Q., and Yang, Y. (2018). Sarcopenia as a predictor of hospitalization among older people: A systematic review and meta-analysis. BMC Geriatr., 18.","DOI":"10.1186\/s12877-018-0878-0"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1154","DOI":"10.1093\/gerona\/glu034","article-title":"Association of sarcopenia with short- and long-term mortality in older adults admitted to acute care wards: Results from the CRIME study","volume":"69","author":"Vetrano","year":"2014","journal-title":"J. Gerontol. Ser. A Biol. Sci. Med. Sci."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"512","DOI":"10.1002\/jcsm.12147","article-title":"Welcome to the ICD-10 code for sarcopenia","volume":"7","author":"Anker","year":"2016","journal-title":"J. Cachexia Sarcopenia Muscle"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"427","DOI":"10.1152\/physrev.00061.2017","article-title":"Sarcopenia: Aging-Related Loss of Muscle Mass and Function","volume":"99","author":"Larsson","year":"2019","journal-title":"Physiol. Rev."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1621","DOI":"10.1096\/fj.13-242750","article-title":"Increased sensitivity to mitochondrial permeability transition and myonuclear translocation of endonuclease G in atrophied muscle of physically active older humans","volume":"28","author":"Gouspillou","year":"2014","journal-title":"FASEB J."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1016\/j.bbabio.2009.09.004","article-title":"Alteration of mitochondrial oxidative phosphorylation in aged skeletal muscle involves modification of adenine nucleotide translocator","volume":"1797","author":"Gouspillou","year":"2010","journal-title":"Biochim. Biophys. Acta (BBA) Bioenerg."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1016\/S0140-6736(89)92143-0","article-title":"Decline in skeletal muscle mitochondrial respiratory chain function: Possible factor in ageing","volume":"1","author":"Trounce","year":"1989","journal-title":"Lancet"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"R519","DOI":"10.1152\/ajpregu.00458.2001","article-title":"Apoptosis in skeletal muscle with aging","volume":"282","author":"Dirks","year":"2002","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1111\/j.1474-9726.2007.00347.x","article-title":"Mitochondrial function and apoptotic susceptibility in aging skeletal muscle","volume":"7","author":"Chabi","year":"2008","journal-title":"Aging Cell"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"5618","DOI":"10.1073\/pnas.0501559102","article-title":"Decline in skeletal muscle mitochondrial function with aging in humans","volume":"102","author":"Short","year":"2005","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1016\/j.cell.2008.06.017","article-title":"Superoxide flashes in single mitochondria","volume":"134","author":"Wang","year":"2008","journal-title":"Cell"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1002\/mus.21526","article-title":"Ros-mediated activation of NF-kappaB and Foxo during muscle disuse","volume":"41","author":"Dodd","year":"2010","journal-title":"Muscle Nerve"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1016\/S0092-8674(04)00400-3","article-title":"Foxo transcription factors induce the atrophy-related ubiquitin ligase atrogin-1 and cause skeletal muscle atrophy","volume":"117","author":"Sandri","year":"2004","journal-title":"Cell"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1089\/rej.2017.1989","article-title":"Circulating Mitochondrial DNA at the Crossroads of Mitochondrial Dysfunction and Inflammation During Aging and Muscle Wasting Disorders","volume":"21","author":"Picca","year":"2018","journal-title":"Rejuvenation Res."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"712","DOI":"10.1038\/s41374-019-0355-1","article-title":"Mitochondrial dysfunction\/NLRP3 inflammasome axis contributes to angiotensin II-induced skeletal muscle wasting via PPAR-gamma","volume":"100","author":"Liu","year":"2020","journal-title":"Lab. Investig. J. Tech. Methods Pathol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"21249","DOI":"10.1074\/jbc.M109.041707","article-title":"Caspase-3 cleaves specific 19 S proteasome subunits in skeletal muscle stimulating proteasome activity","volume":"285","author":"Wang","year":"2010","journal-title":"J. Biol. Chem."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1780","DOI":"10.1152\/jappl.1992.72.5.1780","article-title":"Skeletal muscle adaptations to endurance training in 60- to 70-yr-old men and women","volume":"72","author":"Coggan","year":"1992","journal-title":"J. Appl. Physiol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"517","DOI":"10.1089\/ars.2006.8.517","article-title":"Exercise training attenuates age-induced changes in apoptotic signaling in rat skeletal muscle","volume":"8","author":"Song","year":"2006","journal-title":"Antioxid. Redox Signal."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"St-Jean-Pelletier, F., Pion, C.H., Leduc-Gaudet, J.P., Sgarioto, N., Zovil\u00e9, I., Barbat-Artigas, S., Reynaud, O., Alkaterji, F., Lemieux, F.C., and Grenon, A. (2016). The impact of ageing, physical activity, and pre-frailty on skeletal muscle phenotype, mitochondrial content, and intramyocellular lipids in men. J. Cachexia Sarcopenia Muscle.","DOI":"10.1002\/jcsm.12139"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"721","DOI":"10.2147\/CIA.S132496","article-title":"Nonpharmacological interventions to treat physical frailty and sarcopenia in older patients: A systematic overview\u2014The SENATOR Project ONTOP Series","volume":"12","author":"Abraha","year":"2017","journal-title":"Clin. Interv. Aging"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1093\/gerona\/61.7.675","article-title":"No decline in skeletal muscle oxidative capacity with aging in long-term calorically restricted rats: Effects are independent of mitochondrial DNA integrity","volume":"61","author":"Baker","year":"2006","journal-title":"J. Gerontol. Ser. A Biol. Sci. Med. Sci."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1006\/bbrc.1998.9438","article-title":"Effectiveness of caloric restriction in preventing age-related changes in rat skeletal muscle","volume":"251","author":"Mayhew","year":"1998","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1089\/rej.2006.9.219","article-title":"Caloric restriction protects mitochondrial function with aging in skeletal and cardiac muscles","volume":"9","author":"Hepple","year":"2006","journal-title":"Rejuvenation Res."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1075","DOI":"10.1016\/j.exger.2013.03.004","article-title":"Facts and controversies in our understanding of how caloric restriction impacts the mitochondrion","volume":"48","author":"Gouspillou","year":"2013","journal-title":"Exp. Gerontol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"214","DOI":"10.1002\/(SICI)1097-0185(199706)248:2<214::AID-AR8>3.0.CO;2-S","article-title":"Ultra-High-resolution scanning electron microscopy of mitochondria and sarcoplasmic reticulum arrangement in human red, white, and intermediate muscle fibers","volume":"248","author":"Ogata","year":"1997","journal-title":"Anat. Rec."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1146\/annurev-genet-110410-132529","article-title":"Fusion and fission: Interlinked processes critical for mitochondrial health","volume":"46","author":"Chan","year":"2012","journal-title":"Annu. Rev. Genet."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1577","DOI":"10.1101\/gad.1658508","article-title":"Mitochondrial dynamics and apoptosis","volume":"22","author":"Suen","year":"2008","journal-title":"Genes Dev."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1042\/EBC20170104","article-title":"Mitochondrial dynamics: Overview of molecular mechanisms","volume":"62","author":"Tilokani","year":"2018","journal-title":"Essays Biochem."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"280","DOI":"10.1016\/j.cell.2010.02.026","article-title":"Mitochondrial fusion is required for mtDNA stability in skeletal muscle and tolerance of mtDNA mutations","volume":"141","author":"Chen","year":"2010","journal-title":"Cell"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1677","DOI":"10.15252\/embj.201593084","article-title":"Mfn2 deficiency links age-related sarcopenia and impaired autophagy to activation of an adaptive mitophagy pathway","volume":"35","author":"Sebastian","year":"2016","journal-title":"EMBO J."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1152\/japplphysiol.00719.2018","article-title":"Adult skeletal muscle deletion of Mitofusin 1 and 2 impedes exercise performance and training capacity","volume":"126","author":"Bell","year":"2019","journal-title":"J. Appl. Physiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1374","DOI":"10.1016\/j.cmet.2017.04.021","article-title":"Age-Associated Loss of OPA1 in Muscle Impacts Muscle Mass, Metabolic Homeostasis, Systemic Inflammation, and Epithelial Senescence","volume":"25","author":"Tezze","year":"2017","journal-title":"Cell Metab."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"2126","DOI":"10.15252\/embj.201696179","article-title":"OPA1 deficiency promotes secretion of FGF21 from muscle that prevents obesity and insulin resistance","volume":"36","author":"Pereira","year":"2017","journal-title":"EMBO J."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Rodr\u00edguez-Nuevo, A., D\u00edaz-Ramos, A., Noguera, E., D\u00edaz-S\u00e1ez, F., Duran, X., Mu\u00f1oz, J.P., Romero, M., Plana, N., Sebasti\u00e1n, D., and Tezze, C. (2018). Mitochondrial DNA and TLR9 drive muscle inflammation upon Opa1 deficiency. EMBO J., 37.","DOI":"10.15252\/embj.201796553"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"e1663","DOI":"10.1038\/cddis.2014.595","article-title":"Muscle-specific Drp1 overexpression impairs skeletal muscle growth via translational attenuation","volume":"6","author":"Touvier","year":"2015","journal-title":"Cell Death Dis."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"2383","DOI":"10.1038\/s41418-020-0510-7","article-title":"Drp1 overexpression induces desmin disassembling and drives kinesin-1 activation promoting mitochondrial trafficking in skeletal muscle","volume":"27","author":"Giovarelli","year":"2020","journal-title":"Cell Death Differ."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1774","DOI":"10.1038\/emboj.2010.60","article-title":"Mitochondrial fission and remodelling contributes to muscle atrophy","volume":"29","author":"Romanello","year":"2010","journal-title":"EMBO J."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"2576","DOI":"10.1038\/s41467-019-10226-9","article-title":"DRP1-mediated mitochondrial shape controls calcium homeostasis and muscle mass","volume":"10","author":"Favaro","year":"2019","journal-title":"Nat. Commun."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Dulac, M., Leduc-Gaudet, J.P., Reynaud, O., Ayoub, M.B., Guerin, A., Finkelchtein, M., Hussain, S.N., and Gouspillou, G. (2020). Drp1 knockdown induces severe muscle atrophy and remodelling, mitochondrial dysfunction, autophagy impairment and denervation. J. Physiol.","DOI":"10.1113\/JP279802"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"2091","DOI":"10.1038\/s41467-021-22058-7","article-title":"Myofibril and mitochondria morphogenesis are coordinated by a mechanical feedback mechanism in muscle","volume":"12","author":"Avellaneda","year":"2021","journal-title":"Nat. Commun."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"e13379","DOI":"10.1111\/acel.13379","article-title":"Loss of Fis1 impairs proteostasis during skeletal muscle aging in Drosophila","volume":"20","author":"Lee","year":"2021","journal-title":"Aging Cell"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Joseph, A.M., Adhihetty, P.J., Wawrzyniak, N.R., Wohlgemuth, S.E., Picca, A., Kujoth, G.C., Prolla, T.A., and Leeuwenburgh, C. (2013). Dysregulation of mitochondrial quality control processes contribute to sarcopenia in a mouse model of premature aging. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0069327"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"C422","DOI":"10.1152\/ajpcell.00240.2012","article-title":"Adaptive plasticity of autophagic proteins to denervation in aging skeletal muscle","volume":"304","author":"Vainshtein","year":"2013","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"5509","DOI":"10.1002\/pmic.200900472","article-title":"Proteomic DIGE analysis of the mitochondria-enriched fraction from aged rat skeletal muscle","volume":"9","author":"Ohlendieck","year":"2009","journal-title":"Proteomics"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"963","DOI":"10.1002\/mus.23838","article-title":"Expression of mitochondrial fission and fusion regulatory proteins in skeletal muscle during chronic use and disuse","volume":"48","author":"Iqbal","year":"2013","journal-title":"Muscle Nerve"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"420","DOI":"10.3389\/fphys.2019.00420","article-title":"Effects of Aging and Caloric Restriction on Fiber Type Composition, Mitochondrial Morphology and Dynamics in Rat Oxidative and Glycolytic Muscles","volume":"10","author":"Faitg","year":"2019","journal-title":"Front. Physiol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"R127","DOI":"10.1152\/ajpregu.00337.2011","article-title":"Age-Associated declines in mitochondrial biogenesis and protein quality control factors are minimized by exercise training","volume":"303","author":"Koltai","year":"2012","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"17923","DOI":"10.18632\/oncotarget.4235","article-title":"Mitochondrial morphology is altered in atrophied skeletal muscle of aged mice","volume":"6","author":"Picard","year":"2015","journal-title":"Oncotarget"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1002\/pmic.201500395","article-title":"Specific protein changes contribute to the differential muscle mass loss during ageing","volume":"16","author":"Capitanio","year":"2016","journal-title":"Proteomics"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1016\/j.freeradbiomed.2018.10.456","article-title":"Intensified mitophagy in skeletal muscle with aging is downregulated by PGC-1alpha overexpression in vivo","volume":"130","author":"Yeo","year":"2019","journal-title":"Free Radic. Biol. Med."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1016\/j.exger.2014.02.001","article-title":"Evidence for association of mitochondrial metabolism alteration with lipid accumulation in aging rats","volume":"56","author":"Zhao","year":"2014","journal-title":"Exp. Gerontol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1093\/gerona\/glt107","article-title":"Markers of Human Skeletal Muscle Mitochondrial Biogenesis and Quality Control: Effects of Age and Aerobic Exercise Training","volume":"69","author":"Konopka","year":"2013","journal-title":"J. Gerontol. Ser. A"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1088","DOI":"10.3389\/fphys.2019.01088","article-title":"Regular Endurance Exercise Promotes Fission, Mitophagy, and Oxidative Phosphorylation in Human Skeletal Muscle Independently of Age","volume":"10","author":"Balan","year":"2019","journal-title":"Front. Physiol."},{"key":"ref_58","first-page":"535","article-title":"Chronological Age Does not Influence Ex-vivo Mitochondrial Respiration and Quality Control in Skeletal Muscle","volume":"72","author":"Distefano","year":"2017","journal-title":"J. Gerontol. Ser. A Biol. Sci. Med. Sci."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1111\/j.1474-9726.2012.00844.x","article-title":"The impact of aging on mitochondrial function and biogenesis pathways in skeletal muscle of sedentary high- and low-functioning elderly individuals","volume":"11","author":"Joseph","year":"2012","journal-title":"Aging Cell"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1093\/gerona\/glp179","article-title":"The effect of aging on human skeletal muscle mitochondrial and intramyocellular lipid ultrastructure","volume":"65","author":"Crane","year":"2010","journal-title":"J. Gerontol. Ser. A Biol. Sci. Med. Sci."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"3345","DOI":"10.1113\/JP273950","article-title":"Preservation of skeletal muscle mitochondrial content in older adults: Relationship between mitochondria, fibre type and high-intensity exercise training","volume":"595","author":"Wyckelsma","year":"2017","journal-title":"J. Physiol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1002\/dvg.22050","article-title":"Mouse lines with photo-activatable mitochondria to study mitochondrial dynamics","volume":"50","author":"Pham","year":"2012","journal-title":"Genesis"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"4795","DOI":"10.1113\/jphysiol.2010.199448","article-title":"Repeated transient mRNA bursts precede increases in transcriptional and mitochondrial proteins during training in human skeletal muscle","volume":"588","author":"Perry","year":"2010","journal-title":"J. Physiol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"959","DOI":"10.1096\/fj.15-276907","article-title":"Training intensity modulates changes in PGC-1\u03b1 and p53 protein content and mitochondrial respiration, but not markers of mitochondrial content in human skeletal muscle","volume":"30","author":"Granata","year":"2016","journal-title":"FASEB J."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Meinild Lundby, A.K., Jacobs, R.A., Gehrig, S., de Leur, J., Hauser, M., Bonne, T.C., Fl\u00fcck, D., Dandanell, S., Kirk, N., and Kaech, A. (2018). Exercise training increases skeletal muscle mitochondrial volume density by enlargement of existing mitochondria and not de novo biogenesis. Acta Physiol., 222.","DOI":"10.1111\/apha.12905"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"1033","DOI":"10.1016\/j.cmet.2015.09.027","article-title":"Mitochondrial Dynamics Is a Distinguishing Feature of Skeletal Muscle Fiber Types and Regulates Organellar Compartmentalization","volume":"22","author":"Mishra","year":"2015","journal-title":"Cell Metab."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1111\/apha.12289","article-title":"Three-Dimensional reconstruction of the human skeletal muscle mitochondrial network as a tool to assess mitochondrial content and structural organization","volume":"213","author":"Dahl","year":"2015","journal-title":"Acta Physiol."},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Lally, J.S.V., Herbst, E.A.F., Matravadia, S., Maher, A.C., Perry, C.G.R., Ventura-Clapier, R., and Holloway, G.P. (2013). Over-Expressing Mitofusin-2 in Healthy Mature Mammalian Skeletal Muscle Does Not Alter Mitochondrial Bioenergetics. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0055660"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"448","DOI":"10.1038\/s41467-017-00525-4","article-title":"Promoting Drp1-mediated mitochondrial fission in midlife prolongs healthy lifespan of Drosophila melanogaster","volume":"8","author":"Rana","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1038\/sj.emboj.7601963","article-title":"Fission and selective fusion govern mitochondrial segregation and elimination by autophagy","volume":"27","author":"Twig","year":"2008","journal-title":"EMBO J."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Dulac, M., Leduc-Gaudet, J.P., Cefis, M., Ayoub, M.B., Reynaud, O., Shams, A., Moamer, A., Nery Ferreira, M.F., Hussain, S.N., and Gouspillou, G. (2021). Regulation of muscle and mitochondrial health by the mitochondrial fission protein Drp1 in aged mice. J. Physiol.","DOI":"10.1113\/JP281752"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1038\/nature09782","article-title":"Autophagy in immunity and inflammation","volume":"469","author":"Levine","year":"2011","journal-title":"Nature"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"1542","DOI":"10.1038\/sj.cdd.4401765","article-title":"Autophagy: Molecular machinery for self-eating","volume":"12","author":"Yorimitsu","year":"2005","journal-title":"Cell Death Differ."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1344","DOI":"10.1126\/science.1193497","article-title":"Autophagy and metabolism","volume":"330","author":"Rabinowitz","year":"2010","journal-title":"Science"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1016\/j.cell.2007.12.018","article-title":"Autophagy in the pathogenesis of disease","volume":"132","author":"Levine","year":"2008","journal-title":"Cell"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"814","DOI":"10.1038\/ncb0910-814","article-title":"Eaten alive: A history of macroautophagy","volume":"12","author":"Yang","year":"2010","journal-title":"Nat. Cell Biol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1016\/0014-5793(93)80398-E","article-title":"Isolation and characterization of autophagy-defective mutants of Saccharomyces cerevisiae","volume":"333","author":"Tsukada","year":"1993","journal-title":"FEBS Lett."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"3888","DOI":"10.1242\/jcs.03172","article-title":"Starvation and ULK1-dependent cycling of mammalian Atg9 between the TGN and endosomes","volume":"119","author":"Young","year":"2006","journal-title":"J. Cell Sci."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"18283","DOI":"10.1074\/jbc.M413957200","article-title":"Endothelial nitric-oxide synthase antisense (NOS3AS) gene encodes an autophagy-related protein (APG9-like2) highly expressed in trophoblast","volume":"280","author":"Yamada","year":"2005","journal-title":"J. Biol. Chem."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"1860","DOI":"10.1091\/mbc.e11-09-0746","article-title":"Dynamic and transient interactions of Atg9 with autophagosomes, but not membrane integration, are required for autophagy","volume":"23","author":"Orsi","year":"2012","journal-title":"Mol. Biol. Cell"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1038\/26506","article-title":"A protein conjugation system essential for autophagy","volume":"395","author":"Mizushima","year":"1998","journal-title":"Nature"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"33889","DOI":"10.1074\/jbc.273.51.33889","article-title":"A new protein conjugation system in human. The counterpart of the yeast Apg12p conjugation system essential for autophagy","volume":"273","author":"Mizushima","year":"1998","journal-title":"J. Biol. Chem."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"3888","DOI":"10.1093\/emboj\/18.14.3888","article-title":"Apg16p is required for the function of the Apg12p-Apg5p conjugate in the yeast autophagy pathway","volume":"18","author":"Mizushima","year":"1999","journal-title":"EMBO J."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"1679","DOI":"10.1242\/jcs.00381","article-title":"Mouse Apg16L, a novel WD-repeat protein, targets to the autophagic isolation membrane with the Apg12-Apg5 conjugate","volume":"116","author":"Mizushima","year":"2003","journal-title":"J. Cell Sci."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"1367","DOI":"10.1091\/mbc.10.5.1367","article-title":"Apg7p\/Cvt2p: A novel protein-activating enzyme essential for autophagy","volume":"10","author":"Tanida","year":"1999","journal-title":"Mol. Biol. Cell"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"5234","DOI":"10.1093\/emboj\/18.19.5234","article-title":"Apg10p, a novel protein-conjugating enzyme essential for autophagy in yeast","volume":"18","author":"Shintani","year":"1999","journal-title":"EMBO J."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"18619","DOI":"10.1074\/jbc.M111889200","article-title":"Formation of the approximately 350-kDa Apg12-Apg5.Apg16 multimeric complex, mediated by Apg16 oligomerization, is essential for autophagy in yeast","volume":"277","author":"Kuma","year":"2002","journal-title":"J. Biol. Chem."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"5720","DOI":"10.1093\/emboj\/19.21.5720","article-title":"LC3, a mammalian homologue of yeast Apg8p, is localized in autophagosome membranes after processing","volume":"19","author":"Kabeya","year":"2000","journal-title":"EMBO J."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"2805","DOI":"10.1242\/jcs.01131","article-title":"LC3, GABARAP and GATE16 localize to autophagosomal membrane depending on form-II formation","volume":"117","author":"Kabeya","year":"2004","journal-title":"J. Cell Sci."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1016\/j.molcel.2014.05.021","article-title":"WIPI2 links LC3 conjugation with PI3P, autophagosome formation, and pathogen clearance by recruiting Atg12-5-16L1","volume":"55","author":"Dooley","year":"2014","journal-title":"Mol. Cell"},{"key":"ref_91","doi-asserted-by":"crossref","unstructured":"Klionsky, D.J., Abdel-Aziz, A.K., Abdelfatah, S., Abdellatif, M., Abdoli, A., Abel, S., Abeliovich, H., Abildgaard, M.H., Abudu, Y.P., and Acevedo-Arozena, A. (2021). Guidelines for the use and interpretation of assays for monitoring autophagy (4th edition). Autophagy, 1\u2013382.","DOI":"10.1080\/15548627.2020.1797280"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"1604","DOI":"10.4161\/auto.25955","article-title":"Autophagic flux and oxidative capacity of skeletal muscles during acute starvation","volume":"9","author":"Mofarrahi","year":"2013","journal-title":"Autophagy"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"929","DOI":"10.4161\/auto.6.7.12785","article-title":"Quantitation of \u201cautophagic flux\u201d in mature skeletal muscle","volume":"6","author":"Ju","year":"2010","journal-title":"Autophagy"},{"key":"ref_94","doi-asserted-by":"crossref","unstructured":"Romanello, V., Scalabrin, M., Albiero, M., Blaauw, B., Scorrano, L., and Sandri, M. (2019). Inhibition of the Fission Machinery Mitigates OPA1 Impairment in Adult Skeletal Muscles. Cells, 8.","DOI":"10.3390\/cells8060597"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"5977","DOI":"10.18632\/aging.102990","article-title":"Nrf2 deficiency promotes the increasing trend of autophagy during aging in skeletal muscle: A potential mechanism for the development of sarcopenia","volume":"12","author":"Huang","year":"2020","journal-title":"Aging"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"630","DOI":"10.1002\/jcsm.12409","article-title":"Fibroblast growth factor 21 controls mitophagy and muscle mass","volume":"10","author":"Oost","year":"2019","journal-title":"J. Cachexia Sarcopenia Muscle"},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"e418","DOI":"10.1038\/cddis.2012.159","article-title":"Autophagy as a new therapeutic target in Duchenne muscular dystrophy","volume":"3","author":"Morisi","year":"2012","journal-title":"Cell Death Dis."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/S0076-6879(08)03602-1","article-title":"Methods for monitoring autophagy using GFP-LC3 transgenic mice","volume":"452","author":"Mizushima","year":"2009","journal-title":"Methods Enzymol."},{"key":"ref_99","doi-asserted-by":"crossref","unstructured":"Moulis, M., and Vindis, C. (2017). Methods for Measuring Autophagy in Mice. Cells, 6.","DOI":"10.3390\/cells6020014"},{"key":"ref_100","doi-asserted-by":"crossref","unstructured":"Martinet, W., Roth, L., and De Meyer, G.R.Y. (2017). Standard Immunohistochemical Assays to Assess Autophagy in Mammalian Tissue. Cells, 6.","DOI":"10.3390\/cells6030017"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"3169","DOI":"10.1073\/pnas.76.7.3169","article-title":"Glucagon-induced autophagy and proteolysis in rat liver: Mediation by selective deprivation of intracellular amino acids","volume":"76","author":"Schworer","year":"1979","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"1509","DOI":"10.1016\/j.celrep.2014.07.061","article-title":"Autophagy impairment in muscle induces neuromuscular junction degeneration and precocious aging","volume":"8","author":"Carnio","year":"2014","journal-title":"Cell Rep."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1038\/s41586-018-0162-7","article-title":"Disruption of the beclin 1-BCL2 autophagy regulatory complex promotes longevity in mice","volume":"558","author":"Fernandez","year":"2018","journal-title":"Nature"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1172\/JCI73946","article-title":"Essential role for autophagy in life span extension","volume":"125","author":"Madeo","year":"2015","journal-title":"J. Clin. Investig."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"1387","DOI":"10.1126\/science.1087782","article-title":"Autophagy genes are essential for dauer development and life-span extension in C. elegans","volume":"301","author":"Melendez","year":"2003","journal-title":"Science"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"507","DOI":"10.1016\/j.cmet.2009.10.008","article-title":"Autophagy is required to maintain muscle mass","volume":"10","author":"Masiero","year":"2009","journal-title":"Cell Metab."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.omtm.2017.08.002","article-title":"Atg5(flox)-Derived Autophagy-Deficient Model of Pompe Disease: Does It Tell the Whole Story?","volume":"7","author":"Lim","year":"2017","journal-title":"Mol. Ther. Methods Clin. Dev."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"425","DOI":"10.18632\/aging.100038","article-title":"Mitochondrial dysfunction and oxidative stress mediate the physiological impairment induced by the disruption of autophagy","volume":"1","author":"Wu","year":"2009","journal-title":"Aging"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"2406","DOI":"10.1056\/NEJMoa1915722","article-title":"Developmental Consequences of Defective ATG7-Mediated Autophagy in Humans","volume":"384","author":"Collier","year":"2021","journal-title":"N. Engl. J. Med."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1002\/jcsm.12045","article-title":"p62\/SQSTM1 but not LC3 is accumulated in sarcopenic muscle of mice","volume":"7","author":"Sakuma","year":"2016","journal-title":"J. Cachexia Sarcopenia Muscle"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1111\/j.1474-9726.2007.00282.x","article-title":"Oxidative stress and mitochondrial impairment can be separated from lipofuscin accumulation in aged human skeletal muscle","volume":"6","author":"Skovbro","year":"2007","journal-title":"Aging Cell"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"1513","DOI":"10.1016\/S0531-5565(02)00095-5","article-title":"Distribution of satellite cells in the human vastus lateralis muscle during aging","volume":"37","author":"Renault","year":"2002","journal-title":"Exp. Gerontol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"330","DOI":"10.4161\/auto.5618","article-title":"Longevity pathways converge on autophagy genes to regulate life span in Caenorhabditis elegans","volume":"4","author":"Toth","year":"2008","journal-title":"Autophagy"},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"1029","DOI":"10.1016\/j.celrep.2019.06.070","article-title":"Upregulation of the Autophagy Adaptor p62\/SQSTM1 Prolongs Health and Lifespan in Middle-Aged Drosophila","volume":"28","author":"Aparicio","year":"2019","journal-title":"Cell Rep."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"813","DOI":"10.1016\/j.cell.2010.10.007","article-title":"FOXO\/4E-BP signaling in Drosophila muscles regulates organism-wide proteostasis during aging","volume":"143","author":"Demontis","year":"2010","journal-title":"Cell"},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"458","DOI":"10.1016\/j.cmet.2007.11.001","article-title":"FoxO3 controls autophagy in skeletal muscle in vivo","volume":"6","author":"Mammucari","year":"2007","journal-title":"Cell Metab."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"472","DOI":"10.1016\/j.cmet.2007.11.004","article-title":"FoxO3 coordinately activates protein degradation by the autophagic\/lysosomal and proteasomal pathways in atrophying muscle cells","volume":"6","author":"Zhao","year":"2007","journal-title":"Cell Metab."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"6670","DOI":"10.1038\/ncomms7670","article-title":"Regulation of autophagy and the ubiquitin\u2013proteasome system by the FoxO transcriptional network during muscle atrophy","volume":"6","author":"Milan","year":"2015","journal-title":"Nat. Commun."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"2300","DOI":"10.1038\/ncomms3300","article-title":"Overexpression of Atg5 in mice activates autophagy and extends lifespan","volume":"4","author":"Pyo","year":"2013","journal-title":"Nat. Commun."},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1038\/nature16187","article-title":"Autophagy maintains stemness by preventing senescence","volume":"529","author":"Perdiguero","year":"2016","journal-title":"Nature"},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"795","DOI":"10.1083\/jcb.200809125","article-title":"Parkin is recruited selectively to impaired mitochondria and promotes their autophagy","volume":"183","author":"Narendra","year":"2008","journal-title":"J. Cell Biol."},{"key":"ref_122","doi-asserted-by":"crossref","unstructured":"Narendra, D.P., Jin, S.M., Tanaka, A., Suen, D.-F., Gautier, C.A., Shen, J., Cookson, M.R., and Youle, R.J. (2010). PINK1 is selectively stabilized on impaired mitochondria to activate Parkin. PLoS Biol., 8.","DOI":"10.1371\/journal.pbio.1000298"},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1083\/jcb.201402104","article-title":"PINK1 phosphorylates ubiquitin to activate Parkin E3 ubiquitin ligase activity","volume":"205","author":"Kane","year":"2014","journal-title":"J. Cell Biol."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"378","DOI":"10.1073\/pnas.0911187107","article-title":"PINK1-dependent recruitment of Parkin to mitochondria in mitophagy","volume":"107","author":"Zhou","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1083\/jcb.200910140","article-title":"PINK1 stabilized by mitochondrial depolarization recruits Parkin to damaged mitochondria and activates latent Parkin for mitophagy","volume":"189","author":"Matsuda","year":"2010","journal-title":"J. Cell Biol."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1111\/j.1471-4159.2011.07318.x","article-title":"Parkin promotes the ubiquitination and degradation of the mitochondrial fusion factor mitofusin 1","volume":"118","author":"Glauser","year":"2011","journal-title":"J. Neurochem."},{"key":"ref_127","doi-asserted-by":"crossref","unstructured":"Poole, A.C., Thomas, R.E., Yu, S., Vincow, E.S., and Pallanck, L. (2010). The mitochondrial fusion-promoting factor mitofusin is a substrate of the PINK1\/parkin pathway. PLoS ONE, 5.","DOI":"10.1371\/journal.pone.0010054"},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"5018","DOI":"10.1073\/pnas.0913485107","article-title":"Drosophila parkin requires PINK1 for mitochondrial translocation and ubiquitinates mitofusin","volume":"107","author":"Ziviani","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1126\/science.1231031","article-title":"PINK1-phosphorylated mitofusin 2 is a Parkin receptor for culling damaged mitochondria","volume":"340","author":"Chen","year":"2013","journal-title":"Science"},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1038\/ncb2012","article-title":"PINK1\/Parkin-mediated mitophagy is dependent on VDAC1 and p62\/SQSTM1","volume":"12","author":"Geisler","year":"2010","journal-title":"Nat. Cell Biol."},{"key":"ref_131","doi-asserted-by":"crossref","unstructured":"Liu, S., Sawada, T., Lee, S., Yu, W., Silverio, G., Alapatt, P., Millan, I., Shen, A., Saxton, W., and Kanao, T. (2012). Parkinson\u2019s disease-associated kinase PINK1 regulates Miro protein level and axonal transport of mitochondria. PLoS Genet., 8.","DOI":"10.1371\/journal.pgen.1002537"},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"867","DOI":"10.1093\/hmg\/ddq526","article-title":"PINK1 cleavage at position A103 by the mitochondrial protease PARL","volume":"20","author":"Deas","year":"2011","journal-title":"Hum. Mol. Genet."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"378","DOI":"10.1038\/embor.2012.14","article-title":"Mitochondrial processing peptidase regulates PINK1 processing, import and Parkin recruitment","volume":"13","author":"Greene","year":"2012","journal-title":"EMBO Rep."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"933","DOI":"10.1083\/jcb.201008084","article-title":"Mitochondrial membrane potential regulates PINK1 import and proteolytic destabilization by PARL","volume":"191","author":"Jin","year":"2010","journal-title":"J. Cell Biol."},{"key":"ref_135","doi-asserted-by":"crossref","unstructured":"Kubli, D.A., Cortez, M.Q., Moyzis, A.G., Najor, R.H., Lee, Y., and Gustafsson, A.B. (2015). PINK1 Is Dispensable for Mitochondrial Recruitment of Parkin and Activation of Mitophagy in Cardiac Myocytes. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0130707"},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"939","DOI":"10.1038\/cdd.2009.16","article-title":"Role of BNIP3 and NIX in cell death, autophagy, and mitophagy","volume":"16","author":"Zhang","year":"2009","journal-title":"Cell Death Differ."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"2570","DOI":"10.1128\/MCB.00166-09","article-title":"Hypoxia-Induced Autophagy Is Mediated through Hypoxia-Inducible Factor Induction of BNIP3 and BNIP3L via Their BH3 Domains","volume":"29","author":"Bellot","year":"2009","journal-title":"Mol. Cell. Biol."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"19094","DOI":"10.1074\/jbc.M111.322933","article-title":"Microtubule-associated protein 1 light chain 3 (LC3) interacts with Bnip3 protein to selectively remove endoplasmic reticulum and mitochondria via autophagy","volume":"287","author":"Hanna","year":"2012","journal-title":"J. Biol. Chem."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"721","DOI":"10.1038\/cdd.2010.146","article-title":"Bnip3 impairs mitochondrial bioenergetics and stimulates mitochondrial turnover","volume":"18","author":"Rikka","year":"2011","journal-title":"Cell Death Differ."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1038\/embor.2009.256","article-title":"Nix is a selective autophagy receptor for mitochondrial clearance","volume":"11","author":"Novak","year":"2010","journal-title":"EMBO Rep."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"E4439","DOI":"10.1073\/pnas.1405752111","article-title":"Optineurin is an autophagy receptor for damaged mitochondria in parkin-mediated mitophagy that is disrupted by an ALS-linked mutation","volume":"111","author":"Wong","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1016\/j.molcel.2015.08.016","article-title":"The PINK1-PARKIN Mitochondrial Ubiquitylation Pathway Drives a Program of OPTN\/NDP52 Recruitment and TBK1 Activation to Promote Mitophagy","volume":"60","author":"Heo","year":"2015","journal-title":"Mol. Cell"},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"1357","DOI":"10.1016\/j.celrep.2018.03.127","article-title":"Mitophagy Directs Muscle-Adipose Crosstalk to Alleviate Dietary Obesity","volume":"23","author":"Fu","year":"2018","journal-title":"Cell Rep."},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"1197","DOI":"10.1038\/ncb2837","article-title":"Cardiolipin externalization to the outer mitochondrial membrane acts as an elimination signal for mitophagy in neuronal cells","volume":"15","author":"Chu","year":"2013","journal-title":"Nat. Cell Biol."},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"e13140","DOI":"10.1111\/acel.13140","article-title":"miR-181a regulates p62\/SQSTM1, parkin, and protein DJ-1 promoting mitochondrial dynamics in skeletal muscle aging","volume":"19","author":"McCormick","year":"2020","journal-title":"Aging Cell"},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"3567","DOI":"10.1113\/JP275998","article-title":"Autophagy and mitophagy flux in young and aged skeletal muscle following chronic contractile activity","volume":"596","author":"Carter","year":"2018","journal-title":"J. Physiol."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"E404","DOI":"10.1152\/ajpendo.00391.2017","article-title":"Parkin is required for exercise-induced mitophagy in muscle: Impact of aging","volume":"315","author":"Chen","year":"2018","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1016\/j.exger.2009.03.006","article-title":"Age-Related changes of cell death pathways in rat extraocular muscle","volume":"44","author":"McMullen","year":"2009","journal-title":"Exp. Gerontol."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"192","DOI":"10.1152\/japplphysiol.00175.2011","article-title":"PKB signaling and atrogene expression in skeletal muscle of aged mice","volume":"111","author":"Gaugler","year":"2011","journal-title":"J. Appl. Physiol."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"1040","DOI":"10.1093\/gerona\/glu004","article-title":"Downregulation of E3 ubiquitin ligases and mitophagy-related genes in skeletal muscle of physically inactive, frail older women: A cross-sectional comparison","volume":"69","author":"Drummond","year":"2014","journal-title":"J. Gerontol. Ser. A Biol. Sci. Med. Sci."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1016\/j.exger.2013.11.018","article-title":"Weakness, SR function and stress in gastrocnemius muscles of aged male rats","volume":"50","author":"Russ","year":"2014","journal-title":"Exp. Gerontol."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"1975","DOI":"10.1113\/JP277157","article-title":"Parkin overexpression protects from ageing-related loss of muscle mass and strength","volume":"597","author":"Reynaud","year":"2019","journal-title":"J. Physiol."},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1186\/s13395-018-0157-y","article-title":"Role of Parkin and endurance training on mitochondrial turnover in skeletal muscle","volume":"8","author":"Chen","year":"2018","journal-title":"Skelet Muscle"},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"2565","DOI":"10.1113\/JP275604","article-title":"Protective role of Parkin in skeletal muscle contractile and mitochondrial function","volume":"596","author":"Gouspillou","year":"2018","journal-title":"J. Physiol."},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"C164","DOI":"10.1152\/ajpcell.00064.2017","article-title":"Loss of Parkin impairs mitochondrial function and leads to muscle atrophy","volume":"315","author":"Peker","year":"2018","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"8638","DOI":"10.1073\/pnas.1216197110","article-title":"Parkin overexpression during aging reduces proteotoxicity, alters mitochondrial dynamics, and extends lifespan","volume":"110","author":"Rana","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"4078","DOI":"10.1073\/pnas.0737556100","article-title":"Mitochondrial pathology and apoptotic muscle degeneration in Drosophila parkin mutants","volume":"100","author":"Greene","year":"2003","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"2183","DOI":"10.1242\/dev.01095","article-title":"Drosophila parkin mutants have decreased mass and cell size and increased sensitivity to oxygen radical stress","volume":"131","author":"Pesah","year":"2004","journal-title":"Development"},{"key":"ref_159","doi-asserted-by":"crossref","unstructured":"Leduc-Gaudet, J.P., Mayaki, D., Reynaud, O., Broering, F.E., Chaffer, T.J., Hussain, S.N.A., and Gouspillou, G. (2020). Parkin Overexpression Attenuates Sepsis-Induced Muscle Wasting. Cells, 9.","DOI":"10.3390\/cells9061454"},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"879","DOI":"10.1038\/nm.4132","article-title":"Urolithin A induces mitophagy and prolongs lifespan in C. elegans and increases muscle function in rodents","volume":"22","author":"Ryu","year":"2016","journal-title":"Nat. Med."},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"595","DOI":"10.1038\/s42255-019-0073-4","article-title":"The mitophagy activator urolithin A is safe and induces a molecular signature of improved mitochondrial and cellular health in humans","volume":"1","author":"Andreux","year":"2019","journal-title":"Nat. Metab."},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"14913","DOI":"10.1074\/jbc.M114.556241","article-title":"Systems-based discovery of tomatidine as a natural small molecule inhibitor of skeletal muscle atrophy","volume":"289","author":"Dyle","year":"2014","journal-title":"J. Biol. Chem."},{"key":"ref_163","doi-asserted-by":"crossref","first-page":"46208","DOI":"10.1038\/srep46208","article-title":"Tomatidine enhances lifespan and healthspan in C. elegans through mitophagy induction via the SKN-1\/Nrf2 pathway","volume":"7","author":"Fang","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"345105","DOI":"10.1155\/2014\/345105","article-title":"Antiaging properties of a grape-derived antioxidant are regulated by mitochondrial balance of fusion and fission leading to mitophagy triggered by a signaling network of Sirt1-Sirt3-Foxo3-PINK1-PARKIN","volume":"2014","author":"Das","year":"2014","journal-title":"Oxidative Med. Cell. Longev."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"101048","DOI":"10.1016\/j.isci.2020.101048","article-title":"A Cell-Based High-Throughput Screening Identified Two Compounds that Enhance PINK1-Parkin Signaling","volume":"23","author":"Inoshita","year":"2020","journal-title":"iScience"},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"1034","DOI":"10.1016\/j.cmet.2016.05.007","article-title":"Exercise Promotes Healthy Aging of Skeletal Muscle","volume":"23","author":"Cartee","year":"2016","journal-title":"Cell Metab."},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"548","DOI":"10.1038\/s41467-017-00520-9","article-title":"Ampk phosphorylation of Ulk1 is required for targeting of mitochondria to lysosomes in exercise-induced mitophagy","volume":"8","author":"Laker","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"e13651","DOI":"10.14814\/phy2.13651","article-title":"Exercise and exercise training-induced increase in autophagy markers in human skeletal muscle","volume":"6","author":"Brandt","year":"2018","journal-title":"Physiol. Rep."},{"key":"ref_169","doi-asserted-by":"crossref","first-page":"1447","DOI":"10.1002\/jcsm.12611","article-title":"The effect of fasting or calorie restriction on mitophagy induction: A literature review","volume":"11","author":"Mehrabani","year":"2020","journal-title":"J. Cachexia Sarcopenia Muscle"},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"1048","DOI":"10.1016\/j.cmet.2016.06.001","article-title":"Fasting, Circadian Rhythms, and Time-Restricted Feeding in Healthy Lifespan","volume":"23","author":"Longo","year":"2016","journal-title":"Cell Metab."},{"key":"ref_171","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1016\/j.arr.2016.10.005","article-title":"Impact of intermittent fasting on health and disease processes","volume":"39","author":"Mattson","year":"2017","journal-title":"Ageing Res. Rev."},{"key":"ref_172","doi-asserted-by":"crossref","first-page":"8741","DOI":"10.18632\/aging.102384","article-title":"Time-Restricted eating and age-related muscle loss","volume":"11","author":"Tinsley","year":"2019","journal-title":"Aging"},{"key":"ref_173","doi-asserted-by":"crossref","first-page":"e13322","DOI":"10.1111\/acel.13322","article-title":"The ketogenic diet preserves skeletal muscle with aging in mice","volume":"20","author":"Wallace","year":"2021","journal-title":"Aging Cell"},{"key":"ref_174","doi-asserted-by":"crossref","first-page":"7914","DOI":"10.18632\/aging.202834","article-title":"A ketogenic diet impacts markers of mitochondrial mass in a tissue specific manner in aged mice","volume":"13","author":"Zhou","year":"2021","journal-title":"Aging"},{"key":"ref_175","doi-asserted-by":"crossref","first-page":"382","DOI":"10.3389\/fphys.2019.00382","article-title":"Mitochondrial Quality Control in Aging and Heart Failure: Influence of Ketone Bodies and Mitofusin-Stabilizing Peptides","volume":"10","author":"Thai","year":"2019","journal-title":"Front. Physiol."},{"key":"ref_176","doi-asserted-by":"crossref","unstructured":"Parker, B.A., Walton, C.M., Carr, S.T., Andrus, J.L., Cheung, E.C.K., Duplisea, M.J., Wilson, E.K., Draney, C., Lathen, D.R., and Kenner, K.B. (2018). beta-Hydroxybutyrate Elicits Favorable Mitochondrial Changes in Skeletal Muscle. Int. J. Mol. Sci., 19.","DOI":"10.3390\/ijms19082247"},{"key":"ref_177","doi-asserted-by":"crossref","first-page":"104506","DOI":"10.1016\/j.yrtph.2019.104506","article-title":"Safety and tolerability of sustained exogenous ketosis using ketone monoester drinks for 28 days in healthy adults","volume":"109","author":"Vansant","year":"2019","journal-title":"Regul. Toxicol. Pharmacol. 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