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CLOCK and BMAL1 regulate MyoD and are necessary for maintenance of skeletal muscle phenotype and function
Open accessAbstract
MyoD , a master regulator of myogenesis, exhibits a circadian rhythm in its mRNA and protein levels, suggesting a possible role in the daily maintenance of muscle phenotype and function. We report that MyoD is a direct target of the circadian transcriptional activators CLOCK and BMAL1, which bind in a rhythmic manner to the core enhancer of the MyoD promoter. Skeletal muscle of Clock Δ19 and Bmal1 −/− mutant mice exhibited ∼30% reductions in normalized maximal force. A similar reduction in force was observed at the single-fiber level. Electron microscopy (EM) showed that the myofilament architecture was disrupted in skeletal muscle of Clock Δ19 , Bmal1 −/− , and MyoD −/− mice. The alteration in myofilament organization was associated with decreased expression of actin, myosins, titin, and several MyoD target genes. EM analysis also demonstrated that muscle from both Clock Δ19 and Bmal1 −/− mice had a 40% reduction in mitochondrial volume. The remaining mitochondria in these mutant mice displayed aberrant morphology and increased uncoupling of respiration. This mitochondrial pathology was not seen in muscle of MyoD −/− mice. We suggest that altered expression of both Pgc-1α and Pgc-1β in Clock Δ19 and Bmal1 −/− mice may underlie this pathology. Taken together, our results demonstrate that disruption of CLOCK or BMAL1 leads to structural and functional alterations at the cellular level in skeletal muscle. The identification of MyoD as a clock-controlled gene provides a mechanism by which the circadian clock may generate a muscle-specific circadian transcriptome in an adaptive role for the daily maintenance of adult skeletal muscle.
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“The functional significance of disrupting the molecular clock has been shown in genetic mouse models (i.e., Clock mutant mice and the Bmal1 KO mouse), which display reduced glucose tolerance, mitochondrial respiratory function, and skeletal muscle contractile function [20,21].”
From Exercise mitigates sleep-loss-induced changes in glucose tolerance, mitochondrial function, sarcoplasmic protein synthesis, and diurnal rhythms (2021) · cites this paper as [21]
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“Considering the negative metabolic consequences, such as reduction in insulin sensitivity and mitochondrial function, attributed to both circadian misalignment and sleep loss [21,67] and the potential benefits of exercise in this context [17,68], further research is warranted.”
From Exercise mitigates sleep-loss-induced changes in glucose tolerance, mitochondrial function, sarcoplasmic protein synthesis, and diurnal rhythms (2021) · cites this paper as [21]
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Self-citationCircadian phenotype is associated with diurnal variation in human exercise performance
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Kiperman, T., Xiong, X., Pangemanan, J., Horne, D., Yechoor, V. K., & Ma, K. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.06.05.730283
Nighttime Light Exposure and Frailty: A Large Sample Cross-Sectional StudyPreprint
Ding, Y., Huo, J., Wang, Y., Niu, D., Chen, H., Touré, K., Zhang, J., Yang, Y., Cui, L., Yang, S., et al. (2026) · Research Square · doi:10.21203/rs.3.rs-9850380/v1
Timing the strain: skeletal muscle clock as regulator of physical activity
Kawano, I., & Asher, G. (2026) · npj Metabolic Health and Disease · doi:10.1038/s44324-026-00121-5
“Earlywhole-bodyclockmutantmodels,including Clock Δ 19 , Bmal1 − / − , and Rev-erb α − / − animals, exhibited reduced muscle mitochondrial content and respiratory capacity, disrupted myo fi lament organization, and decreasedcontractileforce 27,31 .”
Xue, M., Wang, H., Xu, P., Wen, H., He, J., Chen, J., Li, H., & Song, C. (2026) · Aquaculture · doi:10.1016/j.aquaculture.2026.744472
Soto, M. D. C., Wang, C., Kang, E., Martinez, S., Toller, J. M., Vasquez, H. E., Herrera, S. V., & Alviña, K. (2026) · Physiology & Behavior · doi:10.1016/j.physbeh.2026.115485
Lee, S., Sim, Y., Lee, M., & Kim, E. (2026) · Antioxidants · doi:10.3390/antiox15091185
Pancreatic Cancer Disrupts Circadian Patterns of Gene Expression in Cardiorespiratory Muscles
Ducharme, J. B., Schonk, M. M., Gutierrez‐Monreal, M. A., Ferreira, L. F., Esser, K. A., Judge, A. R., & Judge, S. M. (2026) · Journal of Cachexia Sarcopenia and Muscle · doi:10.1002/jcsm.70383
Self-citationXue, T., Zhao, Y., Wang, J., Xu, Y., Gu, Z., & Zheng, J. (2026) · Histochemistry and Cell Biology · doi:10.1007/s00418-026-02543-0
“Accumulating evidence indicates that circadian clock genes participate in regulating the myogenic processes of satellite cell function and skeletal muscle regeneration, playing a pivotal role in muscle homeostasis (Andrews et al. 2010; Chatterjee et al. 2013).”
Wang, S., Zhang, Z., Huang, J., Tong, Y., Wu, C., Kobori, H., Seki, Y., Ma, S., & Suzuki, K. (2026) · Nutrients · doi:10.3390/nu18203384
“In skeletal muscle, the integrity of the internal clock contributes to the regulation of glucose metabolism, muscle function, and systemic inflammation [11], and mice lacking functional CLOCK or BMAL1 show reduced muscle force and reduced mitochondrial volume [12].”
Emerging evidence suggests that circadian disruption and muscle wasting form a self-reinforcing loop
Li, F., & Liu, C. (2026) · Chronobiology International · doi:10.1080/07420528.2026.2743668
Investigating the homologue of the mammalian gene myocyte stress 1 in Drosophila melanogaster
Beaumont, K. L. (2013) · Leicester Research Archive (University of Leicester)
THE EFFECTS OF CIRCADIAN RHYTHM ON INFLAMMATORY RESPONSE AND CYTOKINE LEVELS IN HEPATIC MACROPHAGES
Orcino, C. M. (2014) · The Scholarship East Carolina University's Institutional Repository (East Carolina University)
Banduseela, V. C. (2012) · KTH Publication Database DiVA (KTH Royal Institute of Technology)
Metadata
- Authors
- Andrews, J. L., Zhang, X., McCarthy, J. J., McDearmon, E. L., Hornberger, T. A., Russell, B., Campbell, K. S., Arbogast, S., Reid, M. B., Walker, J. R., Hogenesch, J. B., Takahashi, J. S., & Esser, K. A.
- Affiliations
- University of Illinois Chicago (US)
- University of Kentucky (US)
- Northwestern University (US)
- Genomics Institute of the Novartis Research Foundation (US)
- University of Pennsylvania (US)
- ORCID
- Venue
- Proceedings of the National Academy of Sciences
- Publisher
- National Academy of Sciences
- Published
- 2010-10-18
- Volume
- 107
- Issue
- 44
- Pages
- 19090-19095
- ISSN
- 0027-8424 (ISSN-L)
- 1091-6490
- PMC
- PMC2973897
- OpenAlex
- W2053866174
- Cited by
- 389
- References
- 49
- Citations by year
- 2026: 30 · 2025: 29 · 2024: 31 · 2023: 28 · 2022: 36 · 2021: 26 · 2020: 25 · 2019: 19 · 2018: 27 · 2017: 23 · 2016: 32 · 2015: 20 · 2014: 19 · 2013: 17 · 2012: 15
- FWCI
- 6.39
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Proceedings of the National Academy of Sciences (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- HAL (submittedVersion)
- Funders
- Howard Hughes Medical Institute
- National Institutes of Health (5p50mh074924-05)
- National Heart, Lung, and Blood Institute (HL45721, R01HL045721)
- National Institute of Mental Health (U01MH061915, P50 MH-074924, U01 MH 61915)
- National Institute of Arthritis and Musculoskeletal and Skin Diseases (R03AR053641, R21AR050717, AR050717, R01AR055246, AR-053641)
- Topics
- Life Sciences › Neuroscience › Endocrine and Autonomic Systems › Circadian rhythm and melatonin
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Muscle Physiology and Disorders
- Health Sciences › Medicine › Physiology (medical) › Adipose Tissue and Metabolism
- Keywords
- circadian clock, clocks, Bmal1, MyoD, skeletal muscle, myogenesis, clock-controlled genes, mitochondrial dysfunction, uncoupled respiration, PGC-1α, PGC-1β, muscle force, muscle phenotype
- MeSH terms
- Peroxisome Proliferator-Activated Receptor Gamma Coactivator 1-alpha, Animals, Gene Expression Regulation, Mice, Inbred BALB C, Mitochondria, Muscle, Transcription Factors, Trans-Activators, MyoD Protein, Mice, Knockout, Muscle, Skeletal, Gene Expression Profiling, Muscle Development, Mice, Electron Microscope Tomography, CLOCK Proteins, ARNTL Transcription Factors, Circadian Clocks
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