Carbohydrate ingestion influences skeletal muscle cytokine mRNA and plasma cytokine levels after a 3-h run

2003 ◽  
Vol 94 (5) ◽  
pp. 1917-1925 ◽  
Author(s):  
D. C. Nieman ◽  
J. M. Davis ◽  
D. A. Henson ◽  
J. Walberg-Rankin ◽  
M. Shute ◽  
...  
Keyword(s):  
Gene Expression ◽  
Muscle Glycogen ◽  
Glycogen Content ◽  
Vastus Lateralis ◽  
Maximal Oxygen Consumption ◽  
Vastus Lateralis Muscle ◽  
Cytokine Mrna ◽  
Carbohydrate Ingestion ◽  
Muscle Glycogen Content ◽  
Tnf Α

Sixteen experienced marathoners ran on treadmills for 3 h at ∼70% maximal oxygen consumption (V˙o 2 max) on two occasions while receiving 1 l/h carbohydrate (CHO) or placebo (Pla) beverages. Blood and vastus lateralis muscle biopsy samples were collected before and after exercise. Plasma was analyzed for IL-6, IL-10, IL-1 receptor agonist (IL-1ra), IL-8, cortisol, glucose, and insulin. Muscle was analyzed for glycogen content and relative gene expression of 13 cytokines by using real-time quantitative RT-PCR. Plasma glucose and insulin were higher, and cortisol, IL-6, IL-10, and IL-1ra, but not IL-8, were significantly lower postexercise in CHO vs. Pla. Change in muscle glycogen content did not differ between CHO and Pla ( P = 0.246). Muscle cytokine mRNA content was detected preexercise for seven cytokines in this order (highest to lowest): IL-15, TNF-α, IL-8, IL-1β, IL-12p35, IL-6, and IFN-γ. After subjects ran for 3 h, gene expression above prerun levels was measured for five of these cytokines: IL-1β, IL-6, and IL-8 (large increases), and IL-10 and TNF-α (small increases). The increase in mRNA (fold difference from preexercise) was attenuated in CHO (15.9-fold) compared with Pla (35.2-fold) for IL-6 ( P = 0.071) and IL-8 (CHO, 7.8-fold; Pla, 23.3-fold; P = 0.063). CHO compared with Pla beverage ingestion attenuates the increase in plasma IL-6, IL-10, and IL-1ra and gene expression for IL-6 and IL-8 in athletes running 3 h at 70%V˙o 2 max despite no differences in muscle glycogen content.

Influence of carbohydrate ingestion on immune changes after 2 h of intensive resistance training

2004 ◽  
Vol 96 (4) ◽  
pp. 1292-1298 ◽  
Author(s):  
D. C. Nieman ◽  
J. M. Davis ◽  
V. A. Brown ◽  
D. A. Henson ◽  
C. L. Dumke ◽  
...  
Keyword(s):  
Gene Expression ◽  
Glycogen Content ◽  
Weight Training ◽  
Vastus Lateralis ◽  
Vastus Lateralis Muscle ◽  
Carbohydrate Ingestion ◽  
Trained Subjects ◽  
Muscle Gene Expression ◽  
Cell Counts ◽  
Tnf Α

Thirty strength-trained subjects were randomized to carbohydrate (CHO) or placebo (Pla) groups and lifted weights for 2 h (10 exercises, 4 sets each, 10 repetitions, with 2- to 3-min rest intervals). Subjects received 10 ml·kg-1·h-1 CHO (6%) or Pla beverages during the weight training bout. Blood, saliva, and vastus lateralis muscle biopsy samples were collected before and after exercise. Blood cell counts were determined, and plasma was analyzed for IL-6, IL-10, IL-1 receptor antagonist (IL-1ra), IL-8, and cortisol. Muscle was analyzed for glycogen content and relative gene expression of 13 cytokines (IL-1α, IL-1β, IL-2, IL-4, IL-5, IL-6, IL-8, IL-10, IL-12p35, IL-12p40, IL-15, IFN-γ, TNF-α) by use of real-time quantitative RT-PCR. Significant but modest increases were measured for plasma IL-6, IL-10, IL-1ra, and IL-8, but the pattern of increase did not differ between CHO and Pla groups. The rate of decrease in muscle glycogen content did not differ between CHO and Pla ( P = 0.463). Muscle cytokine mRNA was detected preexercise for IL-1β, IL-6, IL-15, IL-8, and TNF-α, and of these, IL-1β, IL-6, IL-8, and TNF-α were significantly increased after the 2-h weight training bout. The increase in mRNA (fold difference from preexercise) did not differ between CHO and Pla groups. In summary, CHO vs. Pla ingestion did not alter modest increases measured for plasma IL-6, IL-10, IL-1ra, and IL-8, and muscle gene expression for IL-1β, IL-6, IL-8, and TNF-α in strength-trained subjects lifting weights intensively for 2 h.

Progressive increase in human skeletal muscle AMPKα2 activity and ACC phosphorylation during exercise

2002 ◽  
Vol 282 (3) ◽  
pp. E688-E694 ◽  
Author(s):  
T. J. Stephens ◽  
Z.-P. Chen ◽  
B. J. Canny ◽  
B. J. Michell ◽  
B. E. Kemp ◽  
...  
Keyword(s):  
Skeletal Muscle ◽  
Muscle Glycogen ◽  
Human Skeletal Muscle ◽  
Glycogen Content ◽  
Vastus Lateralis ◽  
Moderate Intensity ◽  
Moderate Intensity Exercise ◽  
Western Blots ◽  
Muscle Glycogen Content ◽  
Muscle Creatine

The effect of prolonged moderate-intensity exercise on human skeletal muscle AMP-activated protein kinase (AMPK)α1 and -α2 activity and acetyl-CoA carboxylase (ACCβ) and neuronal nitric oxide synthase (nNOSμ) phosphorylation was investigated. Seven active healthy individuals cycled for 30 min at a workload requiring 62.8 ± 1.3% of peak O2consumption (V˙o 2 peak) with muscle biopsies obtained from the vastus lateralis at rest and at 5 and 30 min of exercise. AMPKα1 activity was not altered by exercise; however, AMPKα2 activity was significantly ( P < 0.05) elevated after 5 min (∼2-fold), and further elevated ( P < 0.05) after 30 min (∼3-fold) of exercise. ACCβ phosphorylation was increased ( P < 0.05) after 5 min (∼18-fold compared with rest) and increased ( P< 0.05) further after 30 min of exercise (∼36-fold compared with rest). Increases in AMPKα2 activity were significantly correlated with both increases in ACCβ phosphorylation and reductions in muscle glycogen content. Fat oxidation tended ( P = 0.058) to increase progressively during exercise. Muscle creatine phosphate was lower ( P < 0.05), and muscle creatine, calculated free AMP, and free AMP-to-ATP ratio were higher ( P < 0.05) at both 5 and 30 min of exercise compared with those at rest. At 30 min of exercise, the values of these metabolites were not significantly different from those at 5 min of exercise. Phosphorylation of nNOSμ was variable, and despite the mean doubling with exercise, statistically significance was not achieved ( P = 0.304). Western blots indicated that AMPKα2 was associated with both nNOSμ and ACCβ consistent with them both being substrates of AMPKα2 in vivo. In conclusion, AMPKα2 activity and ACCβ phosphorylation increase progressively during moderate exercise at ∼60% of V˙o 2 peak in humans, with these responses more closely coupled to muscle glycogen content than muscle AMP/ATP ratio.

Effects of cocaine on plasma catecholamine and muscle glycogen concentrations during exercise in the rat

1991 ◽  
Vol 70 (3) ◽  
pp. 1323-1327 ◽  
Author(s):  
R. K. Conlee ◽  
D. W. Barnett ◽  
K. P. Kelly ◽  
D. H. Han
Keyword(s):  
Blood Lactate ◽  
Muscle Glycogen ◽  
Glycogen Content ◽  
Vastus Lateralis ◽  
Male Rats ◽  
Vastus Lateralis Muscle ◽  
Plasma Catecholamine ◽  
Plasma Epinephrine ◽  
Exercise Endurance ◽  
Insight Into

This study was designed to test the hypothesis that cocaine (C) alters the normal physiological responses to exercise. Male rats were injected with saline (S) or C (12.5 mg/kg) either intravenously (iv) or intraperitoneally (ip). After injection the animals were allowed to rest for 30 min or were run on the treadmill (26 m/min, 10% grade). At rest plasma epinephrine values were 245 +/- 24 pg/ml in the S group and 411 +/- 43 (ip) and 612 +/- 41 (iv) pg/ml in the C groups (P less than 0.05 between S and C). During exercise plasma epinephrine levels were 615 +/- 32 pg/ml in S and 1,316 +/- 58 (ip) and 1,208 +/- 37 (iv) pg/ml in the C groups (P less than 0.05 between S and C). Similar results were obtained for norepinephrine. Glycogen content in the white vastus lateralis muscle was reduced to 31 +/- 2 mumol/g in S after exercise, but after C and exercise the values were 12 +/- 4 (ip) and 16 +/- 3 (iv) mumol/g (P less than 0.05 between S and C). There was no effect of the drug on this parameter at rest. Blood lactate rose to 4.8 +/- 1.0 (ip) and 5.8 +/- 1.3 (iv) mM in the C groups but to only 3.0 +/- 0.2 in the S group after exercise (P less than 0.05 between S and C). These results show that C and exercise combined exert a more dramatic effect on plasma catecholamine, muscle glycogen, and blood lactate concentrations than do C and exercise alone. They provide further insight into explaining the adverse effects of C on exercise endurance observed previously (Bracken et al., J. Appl. Physiol. 66: 377-383, 1989).

Influence of muscle glycogen availability on ERK1/2 and Akt signaling after resistance exercise in human skeletal muscle

2005 ◽  
Vol 99 (3) ◽  
pp. 950-956 ◽  
Author(s):  
Andrew Creer ◽  
Philip Gallagher ◽  
Dustin Slivka ◽  
Bozena Jemiolo ◽  
William Fink ◽  
...  
Keyword(s):  
Skeletal Muscle ◽  
Resistance Exercise ◽  
Muscle Glycogen ◽  
Human Skeletal Muscle ◽  
Glycogen Content ◽  
Vastus Lateralis ◽  
Akt Signaling ◽  
Cellular Growth ◽  
Muscle Glycogen Content ◽  
Ribosomal S6 Kinase

Two pathways that have been implicated for cellular growth and development in response to muscle contraction are the extracellular signal-regulated kinase (ERK1/2) and Akt signaling pathways. Although these pathways are readily stimulated after exercise, little is known about how nutritional status may affect stimulation of these pathways in response to resistance exercise in human skeletal muscle. To investigate this, experienced cyclists performed 30 repetitions of knee extension exercise at 70% of one repetition maximum after a low (2%) or high (77%) carbohydrate (LCHO or HCHO) diet, which resulted in low or high (∼174 or ∼591 mmol/kg dry wt) preexercise muscle glycogen content. Muscle biopsies were taken from the vastus lateralis before, ∼20 s after, and 10 min after exercise. ERK1/2 and p90 ribosomal S6 kinase phosphorylation increased ( P ≤ 0.05) 10 min after exercise, regardless of muscle glycogen availability. Akt phosphorylation was elevated ( P < 0.05) 10 min after exercise in the HCHO trial but was unaffected after exercise in the LCHO trial. Mammalian target of rapamycin phosphorylation was similar to that of Akt during each trial; however, change or lack of change was not significant. In conclusion, the ERK1/2 pathway appears to be unaffected by muscle glycogen content. However, muscle glycogen availability appears to contribute to regulation of the Akt pathway, which may influence cellular growth and adaptation in response to resistance exercise in a low-glycogen state.

Effects of eccentric treadmill exercise on inflammatory gene expression in human skeletal muscle

2009 ◽  
Vol 34 (4) ◽  
pp. 745-753 ◽  
Author(s):  
Thomas W. Buford ◽  
Matthew B. Cooke ◽  
Brian D. Shelmadine ◽  
Geoffrey M. Hudson ◽  
Liz Redd ◽  
...  
Keyword(s):  
Gene Expression ◽  
Skeletal Muscle ◽  
Mrna Expression ◽  
Muscle Soreness ◽  
Vastus Lateralis ◽  
Maximal Oxygen Consumption ◽  
Housekeeping Gene ◽  
Necrosis Factor Alpha ◽  
Muscle Gene Expression ◽  
Tnf Α

The present study examined the skeletal muscle expression of several genes related to the inflammatory process before and after a bout of downhill running. Twenty-nine males between the ages of 18 and 35 years performed a 45-min downhill (–17.5%) treadmill protocol at 60% of maximal oxygen consumption. Venous bloods samples and muscle biopsy samples from the vastus lateralis were donated prior to and at 3-h and 24-h postexercise, along with ratings of perceived muscle soreness. Serum creatine kinase (CK) was determined, as was skeletal muscle gene expression of interleukin (IL)-6, IL-8, IL-12 (p35), tumor necrosis factor-alpha (TNF-α), IL-1β, cyclooxygenase 2 (COX2), and nuclear factor kappa B (NFkB) (p105/p50). Gene expression was analyzed using RT-PCR and compared with a standard housekeeping gene (β-actin). Data were analyzed for statistical differences using multivariate analysis of variance with univariate follow-up. In addition, Pearson correlations were conducted to determine if any significant relationship exists between any of these transcripts and both CK and muscle soreness. Significant (p < 0.05) up-regulations in IL-6, IL-8, and COX2 mRNA expression were observed compared with baseline, whereas no significant changes for IL-12, IL-1β, TNF-α, or NFkB were noted. Significant increases in IL-6 mRNA were observed at 3 h (p < 0.001) and 24 h (p = 0.043), whereas significant increases in IL-8 (p = 0.001) and COX2 (p = 0.046) mRNA were observed at 3-h postexercise. In addition, muscle soreness was significantly correlated with IL-8 at 24 h (r = –0.370; p = 0.048), whereas CK was significantly related to NFkB at baseline (r = –0.460; p = 0.012). These data indicate that increases in the mRNA expression of IL-6, IL-8, and COX2 occur in the vastus lateralis as a result of damaging eccentric exercise in young, recreationally trained males. Further, it appears that IL-8 transcription may play some role in inhibiting postexercise muscle soreness, possibly through regulation of angiogenesis.

Muscle glycogen storage after prolonged exercise: effect of the glycemic index of carbohydrate feedings

1993 ◽  
Vol 75 (2) ◽  
pp. 1019-1023 ◽  
Author(s):  
L. M. Burke ◽  
G. R. Collier ◽  
M. Hargreaves
Keyword(s):  
Glycemic Index ◽  
Muscle Glycogen ◽  
Glycogen Content ◽  
Vastus Lateralis ◽  
Glycogen Storage ◽  
Carbohydrate Intake ◽  
Total Carbohydrate ◽  
Muscle Glycogen Content ◽  
Exercise Effect ◽  
Muscle Biopsies

The effect of the glycemic index (GI) of postexercise carbohydrate intake on muscle glycogen storage was investigated. Five well-trained cyclists undertook an exercise trial to deplete muscle glycogen (2 h at 75% of maximal O2 uptake followed by four 30-s sprints) on two occasions, 1 wk apart. For 24 h after each trial, subjects rested and consumed a diet composed exclusively of high-carbohydrate foods, with one trial providing foods with a high GI (HI GI) and the other providing foods with a low GI (LO GI). Total carbohydrate intake over the 24 h was 10 g/kg of body mass, evenly distributed between meals eaten 0, 4, 8, and 21 h postexercise. Blood samples were drawn before exercise, immediately after exercise, immediately before each meal, and 30, 60, and 90 min post-prandially. Muscle biopsies were taken from the vastus lateralis immediately after exercise and after 24 h. When the effects of the immediate postexercise meal were excluded, the totals of the incremental glucose and insulin areas after each meal were greater (P < or = 0.05) for the HI GI meals than for the LO GI meals. The increase in muscle glycogen content after 24 h of recovery was greater (P = 0.02) with the HI GI diet (106 +/- 11.7 mmol/kg wet wt) than with the LO GI diet (71.5 +/- 6.5 mmol/kg). The results suggest that the most rapid increase in muscle glycogen content during the first 24 h of recovery is achieved by consuming foods with a high GI.

Skeletal muscle GLUT-4 and glucose uptake during exercise in humans

1994 ◽  
Vol 77 (3) ◽  
pp. 1565-1568 ◽  
Author(s):  
G. McConell ◽  
M. McCoy ◽  
J. Proietto ◽  
M. Hargreaves
Keyword(s):  
Skeletal Muscle ◽  
Oxygen Consumption ◽  
Glucose Uptake ◽  
Muscle Glycogen ◽  
Protein Level ◽  
Glycogen Content ◽  
Citrate Synthase ◽  
Vastus Lateralis ◽  
Muscle Glycogen Content ◽  
Glut 4

The present study examined the relationship between total skeletal muscle GLUT-4 protein level and glucose uptake during exercise. Eight active non-endurance-trained men cycled at 72 +/- 1% peak pulmonary oxygen consumption for 40 min, with rates of glucose appearance and disappearance (Rd) determined by utilizing a primed continuous infusion of [3–3H]glucose commencing 2 h before exercise. Muscle glycogen content and utilization, citrate synthase activity, and total GLUT-4 protein were measured on muscle biopsy samples obtained from the vastus lateralis. A direct relationship existed between preexercise muscle glycogen content and glycogen utilization during exercise (r = 0.76, P < 0.05). Citrate synthase activity and glucose Rd at the end of exercise averaged 21.9 +/- 3.0 mumol.min-1.g-1 and 27.3 +/- 2.5 mumol.kg-1.min-1, respectively. There was a direct correlation between citrate synthase activity and GLUT-4 protein (r = 0.78, P < 0.05); however, at the end of exercise, glucose Rd was inversely related to both GLUT-4 (r = -0.89, P < 0.01) and citrate synthase activity (r = -0.72, P < 0.05). Plasma insulin, which decreased during exercise, was not related to glucose Rd. In conclusion, glucose uptake during 40 min of exercise at 72% peak pulmonary oxygen consumption was inversely related to the total muscle GLUT-4 protein level. This suggests that factors other than the total GLUT-4 protein level are important in the regulation of glucose uptake during exercise.

scholarly journals The roles of IL-6 in the regulation of glucose homeostasis

2009 ◽  
Vol 34 (1) ◽  
pp. 83-84
Author(s):  
Jenny E. Gusba
Keyword(s):  
Skeletal Muscle ◽  
Protein Kinase ◽  
Muscle Contraction ◽  
Glucose Transport ◽  
Glucose Homeostasis ◽  
Muscle Glycogen ◽  
Glycogen Content ◽  
Glycogen Resynthesis ◽  
Muscle Glycogen Content ◽  
Tnf Α

This thesis examined the roles of interleukin (IL)-6 in the regulation of glucose homeostasis, with a specific focus on skeletal muscle. Study 1 sought to determine whether muscle glycogen content is a stimulus for the production of IL-6, examining the periods during and after exercise. The relationship between IL-6 and muscle glycogen content was measured during similar bouts of exhaustive exercise on 2 occasions that resulted in large increases in muscle messenger (m)RNA for IL-6 and circulating levels of IL-6. On 1 occasion, subjects received carbohydrate during recovery to facilitate rates of glycogen resynthesis. During exercise, subjects performed similar bouts of exercise, such that differences in an individual’s glycogen levels between trials could be compared with differences in IL-6. No correlation was detected between the net change in glycogen content and the net change in plasma IL-6 or IL-6 mRNA from rest to exhaustion. Moreover, when the difference within subjects at exhaustion in IL-6 and glycogen was compared, there was no correlation between the 2 variables. During recovery, although carbohydrate intake significantly increased glycogen resynthesis, there was no change in postexercise IL-6 mRNA level or plasma IL-6 concentration. Therefore, glycogen was not the sole regulator of IL-6 production in skeletal muscle. Study 2 examined the direct effect of IL-6 and tumor necrosis factor (TNF)-α on glucose transport and the phosphorylation of key signalling proteins with or without insulin and during rodent muscle contraction. Under basal conditions, IL-6 increased glucose transport in association with an increase in 5′AMP-activated protein kinase (AMPK) and AS160 phosphorylation, but IL-6 decreased insulin-stimulated glucose transport via a reduction in phosphorylation of calcium–calmodulin-dependent protein kinase (CaMK)II and AS160. A novel finding generated from these experiments was the direct involvement of IL-6 in contraction-mediated glucose transport. In the case of muscle contraction, IL-6 was found to increase the phosphorylation of CaMKII and AS160. This research suggests that the activation of CaMKII is involved in the actions of IL-6 under insulin-stimulated and contraction-mediated conditions. Furthermore, AS160 was identified as a common signalling intermediate, influenced by IL-6. It also suggests that AS160 may be a point of convergence for multiple signalling pathways. Finally, the actions of TNF-α mimicked those of IL-6, except during contraction, where TNF-α had no significant effect on glucose transport and attenuated the effects of IL-6.

scholarly journals Initiating aerobic exercise with low glycogen content reduces markers of myogenesis but not mTORC1 signaling

2021 ◽  
Vol 18 (1) ◽  
Author(s):  
Lee M. Margolis ◽  
Marques A. Wilson ◽  
Claire C. Whitney ◽  
Christopher T. Carrigan ◽  
Nancy E. Murphy ◽  
...  
Keyword(s):  
Aerobic Exercise ◽  
Muscle Glycogen ◽  
Glycogen Content ◽  
Vastus Lateralis ◽  
Cycle Ergometry ◽  
Glycogen Depletion ◽  
Muscle Recovery ◽  
Carbohydrate Ingestion ◽  
Post Exercise ◽  
Mtorc1 Signaling

Abstract Background The effects of low muscle glycogen on molecular markers of protein synthesis and myogenesis before and during aerobic exercise with carbohydrate ingestion is unclear. The purpose of this study was to determine the effects of initiating aerobic exercise with low muscle glycogen on mTORC1 signaling and markers of myogenesis. Methods Eleven men completed two cycle ergometry glycogen depletion trials separated by 7-d, followed by randomized isocaloric refeeding for 24-h to elicit low (LOW; 1.5 g/kg carbohydrate, 3.0 g/kg fat) or adequate (AD; 6.0 g/kg carbohydrate, 1.0 g/kg fat) glycogen. Participants then performed 80-min of cycle ergometry (64 ± 3% VO2peak) while ingesting 146 g carbohydrate. mTORC1 signaling (Western blotting) and gene transcription (RT-qPCR) were determined from vastus lateralis biopsies before glycogen depletion (baseline, BASE), and before (PRE) and after (POST) exercise. Results Regardless of treatment, p-mTORC1Ser2448, p-p70S6KSer424/421, and p-rpS6Ser235/236 were higher (P < 0.05) POST compared to PRE and BASE. PAX7 and MYOGENIN were lower (P < 0.05) in LOW compared to AD, regardless of time, while MYOD was lower (P < 0.05) in LOW compared to AD at PRE, but not different at POST. Conclusion Initiating aerobic exercise with low muscle glycogen does not affect mTORC1 signaling, yet reductions in gene expression of myogenic regulatory factors suggest that muscle recovery from exercise may be reduced.

Fiber type-specific muscle glycogen sparing due to carbohydrate intake before and during exercise

2007 ◽  
Vol 102 (1) ◽  
pp. 183-188 ◽  
Author(s):  
K. De Bock ◽  
W. Derave ◽  
M. Ramaekers ◽  
E. A. Richter ◽  
P. Hespel
Keyword(s):  
Muscle Glycogen ◽  
Glycogen Content ◽  
Fiber Type ◽  
Vastus Lateralis ◽  
Periodic Acid ◽  
Carbohydrate Intake ◽  
Vastus Lateralis Muscle ◽  
Type I ◽  
Type I Fibers ◽  
Glycogen Sparing

The effect of carbohydrate intake before and during exercise on muscle glycogen content was investigated. According to a randomized crossover study design, eight young healthy volunteers ( n = 8) participated in two experimental sessions with an interval of 3 wk. In each session subjects performed 2 h of constant-load bicycle exercise (∼75% maximal oxygen uptake). On one occasion (CHO), they received carbohydrates before (∼150 g) and during (1 g·kg body weight−1·h−1) exercise. On the other occasion they exercised after an overnight fast (F). Fiber type-specific relative glycogen content was determined by periodic acid Schiff staining combined with immunofluorescence in needle biopsies from the vastus lateralis muscle before and immediately after exercise. Preexercise glycogen content was higher in type IIa fibers [9.1 ± 1 × 10−2 optical density (OD)/μm2] than in type I fibers (8.0 ± 1 × 10−2 OD/μm2; P < 0.0001). Type IIa fiber glycogen content decreased during F from 9.6 ± 1 × 10−2 OD/μm2 to 4.5 ± 1 × 10−2 OD/μm2 ( P = 0.001), but it did not significantly change during CHO ( P = 0.29). Conversely, in type I fibers during CHO and F the exercise bout decreased glycogen content to the same degree. We conclude that the combination of carbohydrate intake both before and during moderate- to high-intensity endurance exercise results in glycogen sparing in type IIa muscle fibers.

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