{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T01:07:47Z","timestamp":1790816867988,"version":"4.1.0"},"reference-count":37,"publisher":"MDPI AG","issue":"12","license":[{"start":{"date-parts":[[2020,12,3]],"date-time":"2020-12-03T00:00:00Z","timestamp":1606953600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100002641","name":"Konkuk University","doi-asserted-by":"publisher","award":["KU Research Professor Program"],"award-info":[{"award-number":["KU Research Professor Program"]}],"id":[{"id":"10.13039\/501100002641","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Basic Science Research Program through the National Research Foundation of Korea funded by the Ministry of Science and ICT","award":["2018R1D1A1A02085705"],"award-info":[{"award-number":["2018R1D1A1A02085705"]}]},{"name":"Basic Science Research Program through the National Research Foundation of Korea funded by the Ministry of Science and ICT","award":["2018R1D1A1B07045321"],"award-info":[{"award-number":["2018R1D1A1B07045321"]}]},{"name":"Korea Health Technology R&amp;D Project through the Korea Health Industry Development Institute funded by the Ministry of Health &amp; Welfare","award":["HI18C2383"],"award-info":[{"award-number":["HI18C2383"]}]},{"name":"Korea Basic Science Institute (KBSI) under the R&amp;D programs supervised by the Ministry of Science and ICT","award":["D010710"],"award-info":[{"award-number":["D010710"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Nutrients"],"abstract":"<jats:p>Sarcopenia- or cachexia-related muscle atrophy is due to imbalanced energy metabolism and oxidative stress-induced muscle dysfunction. Monoterpenes play biological and pharmacological reactive oxygen species (ROS) scavenging roles. Hence, we explored the effects of camphene, a bicyclic monoterpene, on skeletal muscle atrophy in vitro and in vivo. We treated L6 myoblast cells with camphene and then examined the ROS-related oxidative stress using Mito TrackerTM Red FM and anti-8-oxoguanine antibody staining. To investigate lipid metabolism, we performed real-time polymerase chain reactions, holotomographic microscopy, and respiratory gas analysis. Rat muscle atrophy in in vivo models was observed using 18F-fluoro-2-deoxy-D-glucose positron emission tomography\/computed tomography and immunocytochemistry. Camphene reversed the aberrant cell size and muscle morphology of L6 myoblasts under starvation and in in vivo models. Camphene also attenuated E3 ubiquitin ligase muscle RING-finger protein-1, mitochondrial fission, and 8-oxoguanine nuclear expression in starved myotubes and hydrogen peroxide (H2O2)-treated cells. Moreover, camphene significantly regulated lipid metabolism in H2O2-treated cells and in vivo models. These findings suggest that camphene may potentially affect skeletal muscle atrophy by regulating oxidative stress and lipid metabolism.<\/jats:p>","DOI":"10.3390\/nu12123731","type":"journal-article","created":{"date-parts":[[2020,12,3]],"date-time":"2020-12-03T11:15:43Z","timestamp":1606994143000},"page":"3731","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":38,"title":["Camphene Attenuates Skeletal Muscle Atrophy by Regulating Oxidative Stress and Lipid Metabolism in Rats"],"prefix":"10.3390","volume":"12","author":[{"given":"Suji","family":"Baek","sequence":"first","affiliation":[{"name":"Research and Development Center, UMUST R&amp;D Corporation, Seoul 05029, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jisu","family":"Kim","sequence":"additional","affiliation":[{"name":"Physical Activity and Performance Institute, Konkuk University, Seoul 05029, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Byung Seok","family":"Moon","sequence":"additional","affiliation":[{"name":"Department of Nuclear Medicine, Ewha Womans University College of Medicine, Seoul 07804, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8546-2006","authenticated-orcid":false,"given":"Sun Mi","family":"Park","sequence":"additional","affiliation":[{"name":"Department of Nuclear Medicine, Ewha Womans University College of Medicine, Seoul 07804, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Da Eun","family":"Jung","sequence":"additional","affiliation":[{"name":"Department of Nuclear Medicine, Ewha Womans University College of Medicine, Seoul 07804, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Seo Young","family":"Kang","sequence":"additional","affiliation":[{"name":"Department of Nuclear Medicine, Ewha Womans University College of Medicine, Seoul 07804, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sang Ju","family":"Lee","sequence":"additional","affiliation":[{"name":"Department of Nuclear Medicine, Asan Medical Center, University of Ulsan College of Medicine, Seoul 05505, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Seung Jun","family":"Oh","sequence":"additional","affiliation":[{"name":"Department of Nuclear Medicine, Asan Medical Center, University of Ulsan College of Medicine, Seoul 05505, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Seung Hae","family":"Kwon","sequence":"additional","affiliation":[{"name":"Seoul Center, Korean Basic Science Institute, Seoul 02841, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Myung Hee","family":"Nam","sequence":"additional","affiliation":[{"name":"Seoul Center, Korean Basic Science Institute, Seoul 02841, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9983-7438","authenticated-orcid":false,"given":"Hye Ok","family":"Kim","sequence":"additional","affiliation":[{"name":"Department of Nuclear Medicine, Ewha Womans University College of Medicine, Seoul 07804, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8752-6280","authenticated-orcid":false,"given":"Hai Jeon","family":"Yoon","sequence":"additional","affiliation":[{"name":"Department of Nuclear Medicine, Ewha Womans University College of Medicine, Seoul 07804, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2520-7182","authenticated-orcid":false,"given":"Bom Sahn","family":"Kim","sequence":"additional","affiliation":[{"name":"Department of Nuclear Medicine, Ewha Womans University College of Medicine, Seoul 07804, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kang Pa","family":"Lee","sequence":"additional","affiliation":[{"name":"Research and Development Center, UMUST R&amp;D Corporation, Seoul 05029, Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2020,12,3]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"2307","DOI":"10.1249\/MSS.0000000000000975","article-title":"Disease-induced skeletal muscle atrophy and fatigue","volume":"48","author":"Powers","year":"2016","journal-title":"Med. Sci. Sports Exerc."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"907","DOI":"10.1016\/j.cell.2004.12.007","article-title":"Molecular mechanisms of muscle atrophy","volume":"119","author":"Mckinnell","year":"2004","journal-title":"Cell"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"434","DOI":"10.3389\/fphys.2017.00434","article-title":"Targeting inflammation and downstream protein metabolism in sarcopenia: A brief updated description of concurrent exercise and leucine-based multimodal intervention","volume":"8","author":"Xia","year":"2017","journal-title":"Front. Physiol."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Fukushima, H., Fujii, Y., and Koga, F. (2019). Metabolic and molecular basis of sarcopenia: Implications in the management of urothelial carcinoma. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20030760"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"240","DOI":"10.1097\/MCO.0b013e328352b4c2","article-title":"Oxidative stress and disuse muscle atrophy: Cause or consequence?","volume":"15","author":"Powers","year":"2012","journal-title":"Curr. Opin. Clin. Nutr. Metab. Care"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"33944","DOI":"10.1038\/srep33944","article-title":"Mitochondrial ROS regulate oxidative damage and mitophagy but not age-related muscle fiber atrophy","volume":"6","author":"Sakellariou","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1007\/s10059-011-0276-3","article-title":"Regulation of reactive oxygen species generation in cell signaling","volume":"32","author":"Bae","year":"2011","journal-title":"Mol. Cells"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"877","DOI":"10.1161\/CIRCRESAHA.117.311401","article-title":"Reactive oxygen species in metabolic and inflammatory signaling","volume":"122","author":"Forrester","year":"2018","journal-title":"Circ. Res."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"208","DOI":"10.1016\/j.freeradbiomed.2016.02.021","article-title":"Redox control of skeletal muscle atrophy","volume":"98","author":"Powers","year":"2016","journal-title":"Free Radic. Biol. Med."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"E469","DOI":"10.1152\/ajpendo.00204.2014","article-title":"Skeletal muscle atrophy and the E3 ubiquitin ligases MuRF1 and MAFbx\/atrogin-1","volume":"307","author":"Bodine","year":"2014","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"158574","DOI":"10.1016\/j.bbalip.2019.158574","article-title":"Lipid accumulation and mitochondrial abnormalities are associated with fiber atrophy in the skeletal muscle of rats with collagen-induced arthritis","volume":"1865","author":"Vial","year":"2020","journal-title":"Biochim. Biophys. Acta Mol. Cell Biol. Lipids"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"993","DOI":"10.1152\/physrev.00038.2012","article-title":"Exercise, GLUT4, and skeletal muscle glucose uptake","volume":"93","author":"Richter","year":"2013","journal-title":"Physiol. Rev."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"633","DOI":"10.1111\/j.1469-7793.1999.0633s.x","article-title":"Lactate transport in skeletal muscle\u2014Role and regulation of the monocarboxylate transporter","volume":"517","author":"Juel","year":"1999","journal-title":"J. Physiol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1089\/jmf.2019.4651","article-title":"High-fat diet increases fat oxidation and promotes skeletal muscle fatty acid transporter expression in exercise-trained mice","volume":"23","author":"Yun","year":"2020","journal-title":"J. Med. Food."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"2044","DOI":"10.1007\/s00125-012-2554-9","article-title":"Inhibition of acetyl-CoA carboxylase 2 enhances skeletal muscle fatty acid oxidation and improves whole-body glucose homeostasis in db\/db mice","volume":"55","author":"Glund","year":"2012","journal-title":"Diabetologia."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"6468593","DOI":"10.1155\/2018\/6468593","article-title":"Essential oils and their major compounds in the treatment of chronic inflammation: A review of antioxidant potential in preclinical studies and molecular mechanisms","volume":"2018","author":"Fernandes","year":"2018","journal-title":"Oxid. Med. Cell Longev."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Vallianou, I., Peroulis, N., Pantazis, P., and Hadzopoulou-Cladaras, M. (2011). Camphene, a plant-derived monoterpene, reduces plasma cholesterol and triglycerides in hyperlipidemic rats independently of HMG-CoA reductase activity. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0020516"},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Ryu, Y., Lee, D., Jung, S.H., Lee, K.J., Jin, H., Kim, S.J., Lee, H.M., Kim, B., and Won, K.J. (2019). Sabinene prevents skeletal muscle atrophy by inhibiting the MAPK-MuRF-1 pathway in rats. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20194955"},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Pasetto, L., Olivari, D., Nardo, G., Trolese, M.C., Bendotti, C., Piccirillo, R., and Bonetto, V. (2018). Micro-computed tomography for non-invasive evaluation of muscle atrophy in mouse models of disease. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0198089"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"250","DOI":"10.3177\/jnsv.59.250","article-title":"Silk peptide intake increases fat oxidation at rest in exercised mice","volume":"59","author":"Kim","year":"2013","journal-title":"J. Nutr. Sci. Vitaminol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1186\/1550-2783-11-35","article-title":"Silk peptide treatment can improve the exercise performance of mice","volume":"11","author":"Kim","year":"2014","journal-title":"J. Int. Soc. Sports Nutr."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1186\/s12986-017-0194-2","article-title":"Piperine enhances carbohydrate\/fat metabolism in skeletal muscle during acute exercise in mice","volume":"14","author":"Kim","year":"2017","journal-title":"Nutr. Metab."},{"key":"ref_23","first-page":"S868","article-title":"The natural compound dansameum reduces foam cell formation by downregulating CD36 and peroxisome proliferator-activated receptor-gamma; expression","volume":"13","author":"Park","year":"2018","journal-title":"Pharmacogn. Mag."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"9397","DOI":"10.1038\/s41598-019-45821-9","article-title":"Skeletal muscle in MuRF1 null mice is not spared in low-gravity conditions, indicating atrophy proceeds by unique mechanisms in space","volume":"9","author":"Cadena","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"e12895","DOI":"10.1111\/acel.12895","article-title":"Reactive oxygen species-induced changes in glucose and lipid metabolism contribute to the accumulation of cholesterol in the liver during aging","volume":"18","author":"Seo","year":"2019","journal-title":"Aging Cell"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1856","DOI":"10.1021\/acsnano.9b07993","article-title":"Label-free tomographic imaging of lipid droplets in foam cells for machine-learning-assisted therapeutic evaluation of targeted nanodrugs","volume":"14","author":"Park","year":"2020","journal-title":"ACS Nano"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"982794","DOI":"10.1155\/2012\/982794","article-title":"Reactive oxygen species in skeletal muscle signaling","volume":"2012","author":"Barbieri","year":"2012","journal-title":"J. Signal Transduct."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"2063179","DOI":"10.1155\/2018\/2063179","article-title":"Role of oxidative stress as key regulator of muscle wasting during cachexia","volume":"2018","author":"Elorza","year":"2018","journal-title":"Oxid. Med. Cell Longev."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1016\/j.tiv.2008.12.014","article-title":"Plant derived antioxidants\u2014Geraniol and camphene protect rat alveolar macrophages against t-BHP induced oxidative stress","volume":"23","author":"Tiwari","year":"2009","journal-title":"Toxicol. In Vitro"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"190","DOI":"10.1002\/jcsm.12144","article-title":"Lipid modulation of skeletal muscle mass and function","volume":"8","author":"Lipina","year":"2016","journal-title":"J. Cachexia Sarcopeni."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s11010-019-03667-9","article-title":"ROS and diseases: Role in metabolism and energy supply","volume":"467","author":"Yang","year":"2020","journal-title":"Mol. Cell Biochem."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1038\/s41419-017-0135-z","article-title":"Cellular death, reactive oxygen species (ROS) and diabetic complications","volume":"9","author":"Volpe","year":"2018","journal-title":"Cell Death Dis."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"941","DOI":"10.1002\/cphy.c100054","article-title":"Reactive oxygen species: Impact on skeletal muscle","volume":"1","author":"Powers","year":"2011","journal-title":"Compr. Physiol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1002\/jcsm.12369","article-title":"Sedentary behaviour, physical activity, and sarcopenia among older adults in the TSHA: Isotemporal substitution model","volume":"10","author":"Ara","year":"2019","journal-title":"J. Cachexia Sarcopeni"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"666","DOI":"10.1038\/nm.4093","article-title":"Excessive fatty acid oxidation induces muscle atrophy in cancer cachexia","volume":"22","author":"Fukawa","year":"2016","journal-title":"Nat. Med."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"69","DOI":"10.3389\/fendo.2016.00069","article-title":"Fatty infiltration of skeletal muscle: Mechanisms and comparisons with bone marrow adiposity","volume":"7","author":"Hamrick","year":"2016","journal-title":"Front. Endocrinol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"141","DOI":"10.3177\/jnsv.62.141","article-title":"Nutrition supplements to stimulate lipolysis: A review in relation to endurance exercise capacity","volume":"62","author":"Kim","year":"2016","journal-title":"J. Nutr. Sci. Vitaminol."}],"container-title":["Nutrients"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2072-6643\/12\/12\/3731\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T10:41:10Z","timestamp":1760179270000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2072-6643\/12\/12\/3731"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,12,3]]},"references-count":37,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2020,12]]}},"alternative-id":["nu12123731"],"URL":"https:\/\/doi.org\/10.3390\/nu12123731","relation":{},"ISSN":["2072-6643"],"issn-type":[{"value":"2072-6643","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,12,3]]}}}