# New insights into the skeletal muscle circadian clock in ruminants

**Authors:** Qiangjun Wang, Yuxin Chen, Yale Chen, Yinghui Ling, Shuai Gao, Zijun Zhang, Dalong Ren

PMC · DOI: 10.1186/s40104-026-01376-0 · 2026-03-13

## TL;DR

This paper reviews how circadian rhythms in ruminant skeletal muscle affect muscle development and meat production.

## Contribution

It systematically explores how management strategies influence muscle satellite cells through the circadian clock.

## Key findings

- Over 2,300 genes in skeletal muscle show circadian expression and are involved in muscle development.
- Disruption of circadian rhythms can impair satellite cell function and muscle growth.
- Environmental and nutritional factors modulate the circadian clock to influence muscle development.

## Abstract

Circadian rhythms are endogenous oscillations with a period of approximately 24 h. They enable organisms to anticipate and adapt to daily environmental changes, such as light and temperature. As the largest metabolic and motor organ in the body, skeletal muscle plays a decisive role in determining meat production efficiency in ruminants. Skeletal muscle development is largely governed by the proliferation and myogenic differentiation capacity of skeletal muscle satellite cells (SMSCs). More than 2,300 genes in skeletal muscle exhibit circadian oscillatory expression and are extensively involved in myogenesis, transcriptional regulation, and metabolic processes. The rhythmic expression of these genes is modulated by external factors including the photoperiod, feeding behavior, gut microbiota, and physical activity. Disruption of the endogenous circadian timing system can inhibit SMSC proliferation and myogenic differentiation, thereby impairing normal muscle development. Therefore, this review focuses on key management aspects of ruminant production—such as environmental control, nutritional regulation, and exercise management—and systematically elaborates on how these husbandry strategies may influence SMSC fate by modulating the circadian clock, along with the underlying molecular mechanisms.

## Full-text entities

- **Genes:** Myhc (myosin heavy chain, cardiac muscle complex) [NCBI Gene 111671], EZH2 (enhancer of zeste 2 polycomb repressive complex 2 subunit) [NCBI Gene 2146] {aka ENX-1, ENX1, EZH2b, KMT6, KMT6A, WVS}, Ezh2 (enhancer of zeste 2 polycomb repressive complex 2 subunit) [NCBI Gene 14056] {aka Enx-1, Enx1h, KMT6, mKIAA4065}, Rps6kb1 (ribosomal protein S6 kinase B1) [NCBI Gene 72508] {aka 2610318I15Rik, P70S6K1, S6K, S6K-beta-1, S6K1, p70 S6K-alpha}, Tmem176b [NCBI Gene 102177498], PIK3CB (phosphatidylinositol-4,5-bisphosphate 3-kinase catalytic subunit beta) [NCBI Gene 5291] {aka P110BETA, PI3K, PI3KBETA, PIK3C1}, DCTN1 [NCBI Gene 102175305], Myf5 [NCBI Gene 100861252], MYF6 (myogenic factor 6) [NCBI Gene 4618] {aka CNM3, MRF4, bHLHc4, myf-6}, Sirt1 (sirtuin 1) [NCBI Gene 93759] {aka SIR2L1, Sir2, Sir2a, Sir2alpha}, Tcap [NCBI Gene 100861162], Tlr4 (toll-like receptor 4) [NCBI Gene 21898] {aka Lps, Ly87, Ran/M1, Rasl2-8}, cyclin D1 [NCBI Gene 102169650], MAP3K7 (mitogen-activated protein kinase kinase kinase 7) [NCBI Gene 6885] {aka CSCF, FMD2, MEKK7, TAK1, TGF1a}, PPARA (peroxisome proliferator activated receptor alpha) [NCBI Gene 5465] {aka NR1C1, PPAR, PPAR-alpha, PPARalpha, hPPAR}, Myf5 (myogenic factor 5) [NCBI Gene 17877] {aka B130010J22Rik, Myf-5, bHLHc2}, IGF1 (insulin like growth factor 1) [NCBI Gene 3479] {aka IGF, IGF-I, IGFI, MGF}, IGF2 (insulin like growth factor 2) [NCBI Gene 3481] {aka C11orf43, GRDF, IGF-II, PP9974, SRS3}, Aanat (arylalkylamine N-acetyltransferase) [NCBI Gene 11298] {aka AA-NAT, Nat-2, Nat4, Snat}, CLOCK [NCBI Gene 102175105], SIRT1 (sirtuin 1) [NCBI Gene 613629], PPARGC1A (PPARG coactivator 1 alpha) [NCBI Gene 10891] {aka LEM6, PGC-1(alpha), PGC-1alpha, PGC-1v, PGC1, PGC1A}, Bmal1 (basic helix-loop-helix ARNT like 1) [NCBI Gene 11865] {aka Arnt3, Arntl, BMAL1b, MOP3, bHLHe5, bmal1b'}, Dbp (D site albumin promoter binding protein) [NCBI Gene 13170], MYD88 (MYD88 innate immune signal transduction adaptor) [NCBI Gene 4615] {aka IMD68, MYD88D, WM1}, CLOCK (clock circadian regulator) [NCBI Gene 100300400], Igf1 (insulin-like growth factor 1) [NCBI Gene 16000] {aka C730016P09Rik, Igf-1, Igf-I}, Mef2c (myocyte enhancer factor 2C) [NCBI Gene 17260] {aka 5430401D19Rik, 9930028G15Rik, Mef2}, CLOCK (clock circadian regulator) [NCBI Gene 9575] {aka KAT13D, bHLHe8}, Nfkb1 (nuclear factor of kappa light polypeptide gene enhancer in B cells 1, p105) [NCBI Gene 18033] {aka NF-KB1, NF-kappaB, NF-kappaB1, p105, p50, p50/p105}, Pik3r1 (phosphoinositide-3-kinase regulatory subunit 1) [NCBI Gene 18708] {aka PI3K, p50alpha, p55alpha, p85alpha}, Cry2 (cryptochrome circadian regulator 2) [NCBI Gene 12953] {aka D130054K12Rik}, BMAL1 (basic helix-loop-helix ARNT like 1) [NCBI Gene 406] {aka ARNTL, ARNTL1, BMAL1c, JAP3, MOP3, PASD3}, NR3C1 (nuclear receptor subfamily 3 group C member 1) [NCBI Gene 2908] {aka GCCR, GCR, GCRST, GR, GRL}, miR-361 [NCBI Gene 104796576], FGF19 (fibroblast growth factor 19) [NCBI Gene 9965], IGF-2 [NCBI Gene 100861277], HSF1 (heat shock transcription factor 1) [NCBI Gene 3297] {aka HSTF1}, Creb1 (cAMP responsive element binding protein 1) [NCBI Gene 12912] {aka 2310001E10Rik, 3526402H21Rik, Creb, Creb-1}, Hsf1 (heat shock factor 1) [NCBI Gene 15499] {aka HSTF, HSTF 1, Hsf1alpha, Hsf1beta}, Shh (sonic hedgehog) [NCBI Gene 20423] {aka 9530036O11Rik, Dsh, HHG-1, Hhg1, Hx, Hxl3}, Csnk1e (casein kinase 1, epsilon) [NCBI Gene 27373] {aka CK1epsilon, CKIe, KC1epsilon, tau}, Pax3 (paired box 3) [NCBI Gene 18505] {aka Pax-3, Sp, Splchl2, splotch}, Per1 (period circadian clock 1) [NCBI Gene 18626] {aka Hftm, Per, m-rigui, mPer1}, MEF2A (myocyte enhancer factor 2A) [NCBI Gene 4205] {aka ADCAD1, RSRFC4, RSRFC9, mef2}, Mstn (myostatin) [NCBI Gene 17700] {aka Cmpt, Gdf8}, PPARGC1A (PPARG coactivator 1 alpha) [NCBI Gene 338446] {aka PPARGC-1A}, PRKAB1 (protein kinase AMP-activated non-catalytic subunit beta 1) [NCBI Gene 5564] {aka AMPK, HAMPKb}, PAX3 (paired box 3) [NCBI Gene 5077] {aka CDHS, HUP2, PAX-3, WS1, WS3}, PAX7 (paired box 7) [NCBI Gene 5081] {aka CMYO19, CMYP19, HUP1, MYOSCO, PAX7B, RMS2}, PTK2B (protein tyrosine kinase 2 beta) [NCBI Gene 2185] {aka CADTK, CAKB, FADK2, FAK2, PKB, PTK}, Mtor (mechanistic target of rapamycin kinase) [NCBI Gene 56717] {aka 2610315D21Rik, FRAP, FRAP2, Frap1, RAFT1, RAPT1}, Adcyap1 (adenylate cyclase activating polypeptide 1) [NCBI Gene 11516] {aka PACAP}, DCTN2 [NCBI Gene 102187922], MYOG (myogenin) [NCBI Gene 4656] {aka MYF4, bHLHc3, myf-4}, SHH (sonic hedgehog signaling molecule) [NCBI Gene 6469] {aka HHG1, HLP3, HPE3, MCOPCB5, SMMCI, ShhNC}, RPS6KB1 (ribosomal protein S6 kinase B1) [NCBI Gene 6198] {aka PS6K, S6K, S6K-beta-1, S6K1, STK14A, p70 S6KA}, MYOD1 (myogenic differentiation 1) [NCBI Gene 4654] {aka CMYO17, CMYP17, MYF3, MYOD, MYODRIF, PUM}, PAX3 [NCBI Gene 102183925], Nfil3 (nuclear factor, interleukin 3, regulated) [NCBI Gene 18030] {aka E4BP4}
- **Diseases:** hypoxic (MESH:D002534), hypertrophy (MESH:D006984), muscular dystrophy (MESH:D009136), muscle atrophy (MESH:D009133), MRFs (MESH:D005171), muscle hypertrophy (MESH:C536106), muscle injury (MESH:D009135), sarcopenia (MESH:D055948), hypoxia (MESH:D000860), SMSC (MESH:D005207)
- **Chemicals:** resveratrol (MESH:D000077185), ATP (MESH:D000255), LPS (MESH:D008070), SCFAs (MESH:D005232), fat (MESH:D005223), glucose (MESH:D005947), Melatonin (MESH:D008550), 5-HT (MESH:D012701), NADH (MESH:D009243), ROS (MESH:D017382), cyclic adenosine monophosphate (MESH:D000242), carbon (MESH:D002244), Fatty acids (MESH:D005227), glutamate (MESH:D018698), vitamin A (MESH:D014801), Bile acids (MESH:D001647), Amino acids (MESH:D000596), branched-chain amino acid (MESH:D000597), BA (MESH:D001464), leucine (MESH:D007930)
- **Species:** Homo sapiens (human, species) [taxon 9606], Ovis aries (domestic sheep, species) [taxon 9940], Bos taurus (bovine, species) [taxon 9913], Capra hircus (domestic goat, species) [taxon 9925], Mus musculus (house mouse, species) [taxon 10090], Oryctolagus cuniculus (domestic rabbit, species) [taxon 9986], Clostridium butyricum (species) [taxon 1492]
- **Cell lines:** SMSC — Homo sapiens (Human), Transformed cell line (CVCL_VG48)

## Figures

6 figures with captions in the complete paper: https://tomesphere.com/paper/PMC12983946/full.md

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Source: https://tomesphere.com/paper/PMC12983946