# MOTS-c References — Full Citation List with DOIs and PubMed Links > Full MOTS-c reference list: 20 peer-reviewed citations from Cell Metabolism, Nature Communications, iScience, Physiological Reports, and others. DOIs and PubMed URLs included. ## Full Reference List All citations are drawn from the peer-reviewed literature. DOIs and PubMed URLs are provided for independent verification. [1] Lee C, Zeng J, Drew BG, Sallam T, Martin-Montalvo A, Wan J, Kim SJ, Mehta H, Hevener AL, de Cabo R, Cohen P. The mitochondrial-derived peptide MOTS-c promotes metabolic homeostasis and reduces obesity and insulin resistance. Cell Metabolism. 2015;21(3):443-454. DOI: 10.1016/j.cmet.2015.02.009 PMID: 25738459 https://pubmed.ncbi.nlm.nih.gov/25738459/ [2] Reynolds JC, Lai RW, Woodhead JST, Joly JH, Mitchell CJ, Cameron-Smith D, Lu R, Cohen P, Graham NA, Benayoun BA, Merry TL, Lee C. MOTS-c is an exercise-induced mitochondrial-encoded regulator of age-dependent physical decline and muscle homeostasis. Nature Communications. 2021;12(1):773. DOI: 10.1038/s41467-020-20790-0 PMID: 33473109 https://pubmed.ncbi.nlm.nih.gov/33473109/ [3] Reynolds JC, et al. [Human exercise data reported in the same study as [2].] Nature Communications. 2021;12(1):773. PMID: 33473109 https://pubmed.ncbi.nlm.nih.gov/33473109/ [4] Kim KH, Son JM, Benayoun BA, Lee C. The Mitochondrial-Encoded Peptide MOTS-c Translocates to the Nucleus to Regulate Nuclear Gene Expression in Response to Metabolic Stress. Cell Metabolism. 2018;28(3):516-524.e7. DOI: 10.1016/j.cmet.2018.06.008 PMID: 29983246 https://pubmed.ncbi.nlm.nih.gov/29983246/ [5] Kim SJ, Miller B, Mehta HH, Xiao J, Wan J, Arpawong TE, Yen K, Cohen P. The mitochondrial-derived peptide MOTS-c is a regulator of plasma metabolites and enhances insulin sensitivity. Physiological Reports. 2019;7(13):e14171. DOI: 10.14814/phy2.14171 PMID: 31293078 https://pubmed.ncbi.nlm.nih.gov/31293078/ [6] D'Souza RF, Woodhead JST, Hedges CP, et al. Increased expression of the mitochondrial derived peptide, MOTS-c, in skeletal muscle of healthy aging men is associated with myofiber composition. Aging (Albany NY). 2020;12(5):5044-5065. DOI: 10.18632/aging.102944 PMID: 32182209 https://pubmed.ncbi.nlm.nih.gov/32182209/ [7] Du C, Zhang C, Wu W, Liang Y, Wang A, Wu S, Zhao Y, Hou L, Ning Q, Luo X. Circulating MOTS-c levels are decreased in obese male children and adolescents and associated with insulin resistance. Pediatric Diabetes. 2018;19(5):1058-1064. DOI: 10.1111/pedi.12685 PMID: 29691953 https://pubmed.ncbi.nlm.nih.gov/29691953/ [8] Luo YH, Xie L, Li JY, Xie Y, Li MQ, Zhou L. Serum MOTS-C Levels are Decreased in Obese Children and Associated with Vascular Endothelial Function. Diabetes, Metabolic Syndrome and Obesity. 2023;16:1125-1133. DOI: 10.2147/DMSO.S403934 PMID: 37077579 https://pubmed.ncbi.nlm.nih.gov/37077579/ [9] Zempo H, Kim SJ, Fuku N, et al. A pro-diabetogenic mtDNA polymorphism in the mitochondrial-derived peptide, MOTS-c. Aging (Albany NY). 2021;13(2):1801-1817. DOI: 10.18632/aging.202529 PMID: 33468709 https://pubmed.ncbi.nlm.nih.gov/33468709/ [10] Zhong P, Peng J, Hu Y, Zhang J, Shen C. Mitochondrial derived peptide MOTS-c prevents the development of heart failure under pressure overload conditions in mice. Journal of Cellular and Molecular Medicine. 2022;26(21):5672-5683. DOI: 10.1111/jcmm.17551 PMID: 36156853 https://pubmed.ncbi.nlm.nih.gov/36156853/ [11] Pham T, Taberner A, Hickey A, Han JC. Mitochondria-derived peptide MOTS-c restores mitochondrial respiration in type 2 diabetic heart. Frontiers in Physiology. 2025;16:1602271. DOI: 10.3389/fphys.2025.1602271 PMID: 40661667 https://pubmed.ncbi.nlm.nih.gov/40661667/ [12] Kumagai H, Kim SJ, Miller B, et al. MOTS-c modulates skeletal muscle function by directly binding and activating CK2. iScience. 2024;27(11):111212. DOI: 10.1016/j.isci.2024.111212 PMID: 39559755 https://pubmed.ncbi.nlm.nih.gov/39559755/ [13] Elhusseiny R, Ihsan M, Bellefroid T, Farooq A, Racinais S, Deldicque L. Mitochondrial-derived peptides MOTS-c and humanin attenuate dexamethasone-induced atrophy in human skeletal muscle cells. Physiological Reports. 2026;14(4):e70791. DOI: 10.14814/phy2.70791 PMID: 41732124 https://pubmed.ncbi.nlm.nih.gov/41732124/ [14] Gao Y, Wei X, Wei P, Lu H, Zhong L, Tan J, Liu H, Liu Z. MOTS-c Functionally Prevents Metabolic Disorders. Metabolites. 2023;13(1):125. DOI: 10.3390/metabo13010125 PMID: 36677050 https://pubmed.ncbi.nlm.nih.gov/36677050/ [15] Jia H, Zhou LC, Chen YF, et al. Mitochondria-encoded peptide MOTS-c participates in plasma membrane repair by facilitating the translocation of TRIM72 to membrane. Theranostics. 2024;14(14):5480-5497. DOI: 10.7150/thno.100321 PMID: 39310099 https://pubmed.ncbi.nlm.nih.gov/39310099/ [16] Domin R, Pytka M, Ruchala M. MOTS-c Serum Concentration Positively Correlates with Lower-Body Muscle Strength and Is Not Related to Maximal Oxygen Uptake — A Preliminary Study. International Journal of Molecular Sciences. 2023;24(19):14951. DOI: 10.3390/ijms241914951 PMID: 37834399 https://pubmed.ncbi.nlm.nih.gov/37834399/ [17] Wan W, Zhang L, Lin Y, Rao X, Wang X, Hua F, Ying J. Mitochondria-derived peptide MOTS-c: effects and mechanisms related to stress, metabolism and aging. Journal of Translational Medicine. 2023;21(1):36. DOI: 10.1186/s12967-023-03885-2 PMID: 36670507 https://pubmed.ncbi.nlm.nih.gov/36670507/ [18] Cobb LJ, Lee C, Xiao J, Yen K, Wong RG, Nakamura HK, Mehta HH, Gao Q, Ashur C, Huffman DM, Wan J, Muzumdar R, Barzilai N, Cohen P. Naturally occurring mitochondrial-derived peptides are age-dependent regulators of apoptosis, insulin sensitivity, and inflammatory markers. Aging (Albany NY). 2016;8(4):796-809. DOI: 10.18632/aging.100943 PMID: 27070352 https://pubmed.ncbi.nlm.nih.gov/27070352/ [19] Mohtashami Z, Singh MK, Salimiaghdam N, Ozgul M, Kenney MC. MOTS-c, the Most Recent Mitochondrial Derived Peptide in Human Aging and Age-Related Diseases. International Journal of Molecular Sciences. 2022;23(19):11991. DOI: 10.3390/ijms231911991 PMID: 36233287 https://pubmed.ncbi.nlm.nih.gov/36233287/ [20] Mohtashami Z, et al. [Bone metabolism data reported in the same study as [19].] International Journal of Molecular Sciences. 2022;23(19):11991. PMID: 36233287 https://pubmed.ncbi.nlm.nih.gov/36233287/ --- Calibrated from the peer-reviewed record — a machined-instrument digest of the MOTS-c literature, not a clinic, not a vendor, not a prescription.