Table 2.

Bioactive compounds derived from whole grains for the regulation of muscle function.

Compounds	Experimental Model	Effective Dose	Targets	Target Process	Effect	References
γ-oryzanol	Male C57BL/6 mice	14.78 mg/kg/d	PPARδ ERRγ	Upregulate mitochondrial biogenesis and promote fatty acid β-oxidation	Reduce muscle weakness, alleviate inflammation and enhance muscle strength	[121]
	Male Wistar rats	0.5% (w/w)	GLUT4	Promote antioxidant and anti-inflammatory effects	Alleviate insulin resistance in muscle	[122]
β-sitosterol	ICR mice C2C12 cells	35 or 350 μg/kg/d 15 μM	UCP3	Upregulate the activity of complex proteins in the mitochondrial electron transport chain and induce mitochondrial uncoupling	Meet energy demand and promote muscle strength	[126]
	Broilers	100 mg/kg	PGC-1α/TFAM signaling	Upregulate oxidative status and mitochondrial biogenesis	Promote muscle performance	[127]
	L6 cells	20 μM	LKB1/AMPK signaling	Increase ACC phosphorylation and triglyceride metabolism	Promote glucose homeostasis and lipid metabolism	[128]
	Male diabetic rats	20 mg/kg/d	Rab/IRAP/Munc 18 pathway	Upregulate GLUT4 translocation and glycolytic and gluconeogenesis enzymes	Alleviate insulin resistance in muscle	[129]
	Male C57BL/6 mice	200 mg/kg/d	FoxO1 signaling	Alleviate muscle protein loss and inhibit protein degradation	Alleviate muscle atrophy	[157]
Alkylresorcinols	Male C57BL/6 mice	0.4% (w/w)	Pyruvate dehydrogenase kinase 4 (PDK4)	Promote lipid-autophagy-induced lipid metabolism disorder	Inhibit the reduction in muscle size and prevent muscle atrophy	[4]
	Male Wistar rats	400 mg/kg/d	Glutathione peroxidase (GPx)	Inhibit oxidative damage caused by lipid accumulation and increase the expression of lactate dehydrogenase (LDH), alanine transaminase (ALT)	Alleviate myocardial infarction and exert an anti-fatigue effect	[130]
	Male C57BL/6J mice	30 or 150 mg/kg/d	SIRT3/PGC-1α signaling	Increase mitochondrial content and mitochondrial biogenesis	Improve exercise capacity	[131]
Ferulic acid	Male SD rats	0.6 mg/kg/d	PKCε	Inhibit lipid-accumulation-induced inflammation	Alleviate insulin resistance	[44]
	Isolated rat psoas muscle cells from Sprague Dawley male albino rats	15–240 µg/mL	α-glucosidase and α-amylase	Promote fatty acid oxidative decomposition and inhibit carbohydrate and lipid hydrolyzing enzymes	Alleviate oxidative stress and mitigate redox imbalance	[5]
	C2C12 cells	25 μM	PI3K/Cpkc signaling	Promote glucose uptake and glycogen synthesis	Improve insulin resistance	[45]
	Zebrafish	0.06 mg/d	MyoD myogenin	Promote muscle growth	Increase the CSA of muscle fiber and muscle mass	[46]
	Duroc × Landrace × Yorkshire (DLY) weaned piglets	0.05% or 0.45% (w/w)	SIRT1/AMPK/PGC-1α signaling	Improve the activity of SDH and MDH, increase the expression of slow MyHC protein	Increase the proportion of slow-twitch fiber and promote mitochondria function	[47]
	C2C12 cells	0.5 or 1 μM	SIRT1/AMPK signaling	Increase the protein level of slow MyHC and decrease the protein level of fast MyHC	Promote slow oxidative muscle fiber formation and inhibit fast muscle fiber formation and exert an anti-fatigue function	[48]
p-coumaric acid	L6 cells	100 μM	AMPK signaling	Promote the fatty acid β-oxidation	Inhibit lipid-accumulation-induced inflammation in muscle	[53]
	C2C12 cells	0.1 mM	AMPK signaling	Increase expression of myogenin and myoD	Improve myogenic differentiation	[52]
Resveratrol	Male C57 BL/6J mice	50 mg/kg/d	AMPK/ FOXO3 signaling	Improve mitochondrial function	Improve muscle atrophy	[55]
	Male C57BL/6J mice	0.4% (w/w)	AMPK/PGC-1α signaling	Increase the level of muscle regeneration proteins including MyoG, Myf5 and Pax7 and mitochondrial biogenesis	Enhance muscle proliferation, differentiation and regeneration of impaired muscle	[57]
	L6 cells	25 μM	PKA/LKB1/ AMPK pathway	Improve mitochondrial dysfunction and oxidative stress	Increase muscle mass and myofiber size and improve induced muscle atrophy	[56]
	Male C57BL/6J mice	15 mg/kg/d		Increase muscle glycogen synthesis and reduce ROS levels	Reduce insulin resistance and promote lipid metabolism	[58]
	C2C12 cells	50 μM	AKT signaling	Modulate redox levels and glucose absorption	Reduce insulin resistance	[59]
	C2C12 cells	100 μM	AKT/mTOR/FOXO1 signaling	Inhibit the atrophy-related ubiquitin ligase	Improve muscle hypertrophy and muscle atrophy	[60]
	Male Kunming mice	400 mg/kg/d	AdiopR1–AMPK–PGC-1α signaling	Increase the expression of myosin heavy chain (MyHC) 1, MyHC2a and MyHC2x	Improve the transformation from fast- to slow-twitch muscle fibers and exercise performance	[61]
	C2C12 cells	50 μM	miR-22-3p	Increase the activities of lactate dehydrogenase (SDH) and malate dehydrogenase (MDH)	Promote muscle-fiber-type conversion from fast-twitch to slow-twitch muscle fibers and exert an anti-fatigue effect	[62]
	Male ICR mice	25 mg/kg/d		Increase the activities of LDH (lactic dehydrogenase) and creatine kinase (CK)	Improve muscle recovery and inflammation	[63]
Quercetin	Male Balb/c mice	0.5% (w/w)	Adiponectin signaling	Decrease the fast MyHC and MyHC IIb protein expression	Promote muscle-fiber-type transformation from fast-twitch to slow-twitch muscle fibers	[66]
	Male C57BL/6 mice	0.05% (w/w)	HO-1/NRF2 signaling	Decrease inflammatory response and oxidative stress	Reduce obesity-induced muscle atrophy	[68]
	Male C57BL/6 mice	0.2% (w/w)	PGC-1α signaling	Improve mitochondrial biogenesis and oxidative phosphorylation	Alleviate disuse-induced muscle atrophy	[69]
	C2C12 cells	20 µM	AMPK signaling	Enhance insulin-stimulated glucose uptake and decrease inflammatory response	Ameliorate inflammation-induced insulin resistance	[70]
Oligomeric procyanidins (OPCs)	Male ICR mice	15 mg/kg/d	mTOR signaling	Increase glucose uptake and glycolysis, improve heat generation and inhibit gluconeogenesis and lipogenesis	Improve glucose homeostasis, lipid metabolism and insulin sensitivity	[75]
	Human primary skeletal muscle cells	10 or 25 μM	AKT signaling	Increase glycogen synthesis and glucose uptake	Improve glucose utilization and alleviate insulin resistance	[76]
	Male ICR mice	10 μg/kg/d	AMPK signaling	Promote GLUT4 translocation	Increase insulin sensitivity	[77]
Cyanidin-3-glucoside (Cy3G)	Male ICR mice	1 mg/kg/d	PGC-1α signaling	Improve mitochondrial content and mitochondrial biogenesis	Improve exercise capacity	[82]
	Human skeletal muscle cells	100 µM		Inhibit the activity of α-amylase and α-glucosidase	Alleviate diabetes	[81]
Catechin	C2C12 cells	20 µM	MyoD, MyoG, and MyHC	Promote myotube differentiation	Improve skeletal muscle regeneration and repair	[87]
	Male C57BL/6 mice	25 mg/kg/d	Wnt signaling	Promote myotube differentiation	Promote muscle regeneration	[90]
	C2C12 cells	10 μM	Akt	Promote myotube differentiation	Improve muscle regeneration	[86]
	Male rats	1.0 mg/kg/d	p38 MAPK signaling	Promote mitochondrial respiratory capacity and mitochondrial biogenesis	Enhance the ability of resisting fatigue	[88]
	Male Goto–Kakizaki (GK) rats	100 mg/kg/d	ROS-ERK/JNK-p53 pathway	Promote mitochondrial autophagy	Alleviate diabetic-induced sarcopenia	[89]
	Male Sprague Dawley rats	200 mg/kg/d	IGF-1	Downregulate UPS-mediated muscle protein degradation and upregulate IGF-1-mediated muscle protein synthesis	Increase muscle mass	[91]
	Male SD rats	0.1% (w/w)		Inhibit the expression of oxidative-modified proteins	Prevent muscle from oxidative stress induced by free radicals	[92]
Rutin	Male Sprague Dawley rats	0.1% (w/w)	AMPK signaling	Enhance mitochondrial DNA (mtDNA) content and mitochondrial biogenesis	Improve obesity-induced muscle mitochondrial dysfunction	[97]
	Male C57BL/6 mice	60 mg/kg/d	PGC-1α signaling	Upregulate mediated mitochondrial biogenesis and decrease the level of lactic acid	Improve fatigue-resistance capacity	[98]
	C2C12 cells	100 μM	NF-κB signaling	Decrease the expression of IL-6 and iNOS and the production of ROS	Inhibit oxidative-stress-induced skeletal muscle injury	[99]
Lutein	Male Wistar rats	0.5 mg/kg/d	NF-κB signaling	Reduce oxidative stress and inflammation and decrease the production of ROS	Improve skeletal muscle IR injury	[103]
β-carotene	Male Kwl: ddY mice	0.5 mg/kg/d	IGF-1	Promote protein synthesis and reduce ubiquitin-mediated muscle protein degradation	Increase muscle mass and prevent muscle hypertrophy	[105]
	C2C12 cells	10 μM	FOXO3A	Decrease the level of Atrogin-1 and MuRF1	Increase muscle mass and exhibit an improvement effect on oxidative-stress-induced muscle atrophy	[107]
Betaine	C2C12 cells	10 mM	NFATc1/ MyoD signaling	Upregulate the expression of miR-29b-3p and promote myotube differentiation and the expression of slow MyHC proteins	Promote muscle cell differentiation and the transformation from fast muscle to slow muscle fiber	[136]
	C2C12 cells	10 mM	IGF-1 signaling	Increase the expression of MyoD and myogenin	Promote muscle fiber differentiation and growth	[134]
	C2C12 cells	2 or 5 mM	PGC-1α signaling	Increase mitochondrial biogenesis and ATP production	Promote muscle differentiation and the transformation from fast muscle to slow muscle fiber	[137]
Octacosanol	Male Wistar rats	2.0 μCi/dose		Promote energy mobilization and energy supply	Enhance physical performance	[140]
	Male C57BL/6 mice	200 mg/kg/d	Bcl3/TLRs/MAPK signaling	Increase the expression of Prx, Trim63 and ATPase activity	Exert an anti-fatigue effect	[143]
	Male SD rats	0.75% (w/w)	Creatine phosphorylation	Promote the shift from fast-twitch to slow-twitch myofibers	Exert an anti-fatigue effect	[141]
	Male Wistar rats	2.0 μCi/dose	ACC phosphorylation	Meet energy demand	Improve exercise capacity	[142]
β-glucan	Male SD rats	312.5 mg/kg/d		Decrease the activity of lactate dehydrogenase and the creatine kinase	Increase exercise capacity and facilitate the recovery from fatigue	[116]
	C2C12 cells	20 mg/mL	Myf5 and Mox2	Increase muscle cell proliferation and differentiation	Promote the transformation from fast muscle fibers to slow muscle fibers	[117]
	Duchenne muscular dystrophy (DMD) zebrafish model	8 mg/L	Mitochondrial respiration enzyme	Improve mitochondrial respiration and prevent oxidative stress	Improve exercise capacity	[118]
Tocotrienol	Male C57BL/6J mice	100 or 300 mg/kg/d	AMPK/SIRT1/PGC-1α signaling	Upregulate the expression of proliferation and differentiation related proteins Increase mitochondrial biogenesis	Prevent diabetes-related skeletal muscle atrophy	[110]
	Human skeletal muscle myoblasts	50 μg/mL	miR-206	Increase the expression of IGF1R and decrease the expression of Pax7	Promote muscle cell proliferation and differentiation	[111]
	Stress-induced premature senescence (SIPS) Human skeletal muscle myoblasts (CHQ5B)	50 μg/mL	Wnt signaling FOXO pathway	Downregulate the expression of MSTN and increase the expression of muscle cells regeneration related proteins such as EREG, SHC1 and SHC3	Promote muscle cell regeneration and alleviate muscle loss	[112]
	Male C57BL/6J mice	400 mg/kg/d	COXⅠ-Ⅴ	Promote mitochondrial respiration and reduce lipid peroxidation	Increase muscle mass and improve glucose homeostasis	[113]